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Bulletin of the Museum of Comparative Zoology 

AT HARVARD COLLEGE 

Vol, 127, No. 5 



IHU 



ITU i tUUb 



LIBRARY 



I 






JUL 2 3 1962 




UNIVERSITY 



ON SOME BURMESE EARTHWORMS OF THE 
MONILIGASTRID GENUS DBAW1DA 



By G. E. Gates 

University of Maine, Orono 



CAMBRIDGE, MASS., U.S.A. 
PRINTED FOR THE MUSEUM 

July 23, 1962 



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Bulletin of the Museum of Comparative Zoology 

AT HAKVAED COLLEGE 

Vol. 127, No. 5 



ON SOME BURMESE EARTHWORMS OF THE 
MONILIGASTRID GENUS DRAWIDA 



By G. E. Gates 

University of Maine, Orono 



CAMBRIDGE, MASS., U.S.A. 
PRINTED FOR THE MUSEUM 

July, 1962 




1MUS. COMP. ZOOl 
LIBRARY 

JUL 2 3 1962 

HARVARD 
UNIVERSITY 




Bull, Mm. Comp. Zooh, Harvard Uiiiv,, 127(5): 295-374, July, 1962 

No. 5 — On Some Burmese Earthworms of the Moniligastrid 

Genus Drawl da 1 

By G. E. Gates 



CONTENTS 

Introduction 297 

Systematica . . . - . « . . . . 298 

Oligoehaeta, Class or Order . 298 

Family Moniligastridae . . -99 

Genus Drawida Michaelaen, 1900 ... . . 305 

Taxonomie characters 308 

Drawida beddardi (Rosa, 1890) . . .......... 312 

Drawida bidlaia Gates, 1933 313 

Drawida eaerulea Gates, 1926 .,, 317 

Drawida delicata nom, nov, 319 

Drawida flexa Gates, 1929 320 

Drawida gracilis Gates, 1925 324 

Drawida long atria Gates, 1925 326 

Dratvida nana Gates, 1933 330 

Drawida nepalensis Michaelscn, 1907 331 

Drawida papilUfer Stephenson, 1917 . . 340 

Drawida ranffoonensig Gates, 192*1 . , , , 346 

Dratvida vara Gates, 1925 . . 348 

Drawida tenellula nom. nov. . . . , 351 

Drawida vietoriana sp. nov, ... 352 

Drawida vvXgaris Gates, 1930 353 

Genus Momligaster Perrier, 1872 . . 355 

Maniligaster cernosvitovi nom, nov. . . 355 

.Moniligastrid phylogeiiy ... 355 

Oligochaete phytogeny . . ...♦., 362 

References 372 



INTRODUCTION 

This contribution completes publication of results of the au- 
thor's study of Burmese moniligastrids. Specimens listed herein, 
with few exceptions, were collected between 1932 and the end 
of 1941 when the survey was finally terminated by the Japanese 
invasion. Locality lists for the period subsequent to 1932 prob- 
ably are complete for most of the species considered below, 
though information about certain anatomical and other matters 

i From a manuscript written during toimre of a John Simon Guggenheim 
Fellowship, 




298 BULLETIN : MUSEUM OF COMPABATIVE ZOOLOGY 

is available no longer. Some comparable data for one species was 
obtained recently, through assistance of Dr. II. K. Bhatti, from 
a Pakistani series. Unpublished records of other Burmese 
drawidas were destroyed along with the specimens during World 
War II. 

The Burma survey provided, for the first time, considerable 
material of moniligastrid species. The variation, both individual 
and geographic, that was revealed required consideration (p. 
308) of the usefulness to taxonomy of various characters. In 
addition to the taxonomic accounts, discussions of anatomy (pp, 
299 and 305) provide the foundation for a first attempt at a 
phytogeny of the family (p. 355), as well as for discussion of 
certain assumptions basic in classical phytogenies. An iL in- 
genious ?? contraction theory, the only previous attempt to bridge 
the gap between moniligastrids and other megadriles, is critically 
examined for the first time (p. 363), and other ways in which 
early oligochaete evolution may have proceeded are suggested. 

SYSTEMATICS 
OLIGOCHAETA, Class or Order 

The lower rank was acceptable to classical authorities, Bed* 
dard (1895), Michaelscn (1000, 1921, 1928-1930) and Stephen- 
son (1923, 1930), The order, along with the Polychaeta, long 
was included in a class Chaetopoda. Since 1900, oHgochaetes 
have been found to be more closely related to the leeches than 
to the polychaetes. Michaelsen's Acanthobdellidae, in Kiikenthal 
and Krumbach's massive "Handbueh der Zoologie/ 7 is in the 
Oligochaeta and also in the Hirudinea! To show the relationship, 
some authorities, including Stephenson (1930), place the leeches 
also in the Chaetopoda, Michaelsen, however, erected for the 
two more closely related orders the class Clitellata which has not 
been generally accepted. Acantkobdella was retained in the 
Oligochaeta by Piekford (1948) and Yamaguchi (1953) but was 
excluded by Stephenson (1930) and Avel (1959). Though "The 
great gulf in intra-annelidan phylogeny is that which divides 
the polychaetes from the oHgochaetes" Piekford (1948) believed 
that convenience required the three main annelidan groups to 
have equal rank as classes. Tf a classification is to indicate the 
closer leech-oligochaete relationship, some place presumably ought 
to be found for the Clitellata. 

Between order and families, in his later systems Michaelsen 
interposed two (1921), then three (1928), and finally four (1930) 



GATES : BURMESE MONILIGASTRIDS 299 

suborders as well as various series. These taxa were accepted 
by Pickford, Yamaguchi and Avel though with some modifica- 
tions, including raising suborders (Pickford, Avel) to ordinal 
rank. This elaboration of the classification, in agreement with 
Stephenson (1930, p. 719), seems unwarranted by our present 
knowledge, 

Pickford and Avel, as well as Stephenson, inadvertently reveal 
in their discussions a more immediate need for distinguishing 
two groups of oligochaetes. Three sets of terms have been used 
in the past : Terricolae and Limicolae, Megadrili and Mierodrili, 
higher and lower oligochaetes. Many species obviously of earth- 
worm size (sometimes as much as 12-18 mm. thick) and of 
terricolous habit formerly were considered to be mierodrile or 
limicole. By 1930 Stephenson admitted it was sometimes con- 
venient to include moniligastrids in the Megadrili. Later Yama- 
guchi (1953, p. 331) added the Haplotaxidae, Alluroididae and 
Syngenodrilidae. The terms indicative of size no longer are 
inappropriate though a very few microdriles may be somewhat 
larger than smallest megadriles. The Haplotaxidae, Alluroididae 
and Syngenodrilidae as well as the Moniligastridae, for more than 
thirty years, have been lumped together by the writer (ef. Gates, 
1959) as " earthworms". No common word for "non-earth- 
worms," however, has been available. Yamaguchi f s division of 
the oligochaetes, which is in agreement with the author's prac- 
tice, provides an appropriate pair of terms. The Megadrili, 
accordingly, includes the fourteen families of earthworms that 
are now recognized (Gates, 1959). The Mierodrili comprises the 
remaining oligochaete families including the Aeanthobdellidae, 
The megadrile group is the exact equivalent of Yamaguchi f s 
Opisthopora diplotestieulata and of Michaelsen's Opisthopora. 
The latter was defined eircuitously and, like other suborders and 
series, on vast extrapolations from very little data. 

Megadrili and Mierodrili, in the author's usage, are only con- 
venient terms to facilitate communication. Accordingly, they 
require no formal definition or morphological characterization. 

Family MONILIGASTRIDAE 

This megadrile group has been conspicuous among classical 
families because of its homogeneity. Isolation from other oligo- 
chaetes was recognized early and the family could have been bet- 
ter defined as long ago as 1895 from information then available. 
Nevertheless, unique structure enabling recognition of isolation 



300 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

has not yet been mentioned in a formal definition. Diagnostic 
characters are: intraseptal location of testes and male funnels, 
delimitation of an ovarian chamber from peri-esophageal and 
neural coelomic spaces, presence of enterosegmental organs, me- 
dian union of last pair (or each of last two pairs) of hearts above 
the gut to open into dorsal trunk indirectly through a short 
vertical vessel. All of those characters were retained by the 
most aberrant individuals that have been found. In contrast to 
so much uniqueness, other mcgadrile families are defined by no 
single diagnostic character or by just one for a group (Megaseo- 
lecidae) in which distinctive organs are lacking in partheno- 
genetic morphs. 

A moniligastrid character, perhaps less likely to be unique, is 
provided by the prostomium. That organ, presumably as a result 
of considerable deepening and subsequent posterior extension 
of a slight transverse groove that once provided demarcation 
from the peristomium, is now attached to the roof of the buccal 
cavity behind level of intersegmental furrow 1/2. Still pro- 
trusible, and also capable of assuming a sucker-like shape, the 
moniligastrid prostomium at present appears not to be homolo- 
gous with the longer and slenderer proboscis of certain glossoseo- 

lecids* 

Absence of seminal vesicles is not confined to the Moniligas- 
tridae as those organs have disappeared in recent parthenogenetie 
morphs of several families. Extra-esophageal blood vessels are 
also lateral to the hearts in Stfiiff* nodrUtis and may prove to be 
similarly located in other genera when long neglected vascular 
systems are studied. The single -layered clitellum, yolky ova, and 
location of male pores in front of the female apertures are all 
characters supposedly shared with the Alluroididae and Hap- 
lotaxidae as well as microdrile families. 

Each portion of a moniligastrid septum containing a testis and 
male funnel has been called a testis sac. That characterization 
originally was given to, and in most megadriles still refers to, 
a truly coelomic space, containing one or both testes and male 
funnels of a segment, bounded by membranous or muscularized 
partitions, in communication with the exterior by an aperture 
in the funnel and the narrow gonoduct lumen as well as a minute 
male pore. Such a sac often is formed by development of par- 
titions that seal off one (an unpaired sac) or two (paired sacs) 
ventral portions of the coelomic cavity. In various other mega- 
driles the sac results from a series of gradual evolutionary 
changes such as approximation (or apparently so) of parietal 



GATES : BURMESE M0NIL1GASTRIDS 301 

insertions of two consecutive septa, apposition of the septa 
peripherally finally followed occasionally by abortion of dorsal 
portions of the united septa. In all such cases the coelomic 
cavity of a particular segment is more or less markedly reduced 
but without affecting size of the metamere or diminishing total 
volume of coelomic space. Ocnerodrilid testis sacs are of a dif- 
ferent sort. Each is merely a testis much enlarged by retention 
of developing gametes. The sac is a solid protuberance from the 
posterior face of the septum and may reach considerable size, 
even large enough to have been mistaken on various occasions 
for seminal vesicles. Any internal space (not artificially pro- 
duced) that may appear is, of course, schizoeoelic and not coelo- 
mic. The sac, unlike the seminal vesicles, is not in communication 
with the eoelom nor does it open to the exterior. Sperm are 
released into the coelomic cavity by rupture of the sac wall and 
pass to the exterior via gonoducts that open to the interior and 
exterior as usual. The moniligastrid sac likewise is solid, any 
spaces not produced by preservation, fixation or dehydration, 
etc., in preparation for sectioning, again must be sehizoeoelic. 
Differences from the ocnerodrilid sac are two. First, the develop- 
ing germ cells are proliferated, not posteriorly, but into the 
interior of a septum which becomes bulged anteriorly as well as 
posteriorly and more or less equally (if conditions permit) into 
two consecutive coelomic cavities. Second, the sac also contains 
a male funnel on which the testis sometimes even seems to be 
seated. Sperm no longer are released into the eoelom to run the 
risk of ingestion by amoeboeytes. Moniligastrid and ocnerodrilid 
testis sacs obviously are not homologous with any of the various 
kinds found in other megadriles. 

The cavity of the moniligastrid ovarian chamber, unlike any 
schizocoelic spaces that may be present in otherwise solid testis 
sacs, is coelomic. The chamber, in more primitive forms, seems 
to be still bounded by the parietes peripherally but is closed off 
mesially from periesophageal and neural portions of the eoelom 
that are omitted in figures purporting to show similarity to the 
testis sac. The chamber is closed off, in more advanced forms, 
from the parietes and then is arched like a horseshoe over the 
gut with one limb passing ventrally on each side. The ovarian 
segment is just as large as the one next behind, hence there is 
no reduction in size, merely a rearrangement of internal par- 
titions. Complete closure of the chamber, rather curiously, does 
provide at maximal distension much more space for yolk and 
ova than had been available before. 




302 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Anatomy of the enteroseginental organs has been studied only 
in one species, D. nepalrnsis (el. below). Function is unknown 
and the noncommittal name is intended only to indicate that they 
are metamerically repented and are on the gut, into which each 
component may open at one or both ends. 

The esophagus, in most megadriles studied by the author and 
also in South African aeanthrodrilids (Piekford, 1937), ends in 
a narrow valve that must relax to allow passage of ingesta into 
the intestine. As moniligastrid gizzards long had been said to 
be at the beginning of the intestine, a valve was sought, in front 
of the first gizzard but was not found. A narrowed portion of 
the gut much like a valve, concealed from view by deep saccu- 
lations of adjacent portions of the gut, eventually was located 
some distance behind the last gizzard. Thereafter, all of the 
gut between pharynx and the valve, regardless of length, was 
regarded as esophageal. The eutieular lining has been traced 
from the gizzards through several segments but not as yet all 
the way to the valve or to the buccal cavity. Determination of 
the segment of intestinal origin proved to be much more difficult 
than in other megadriles, partly because of close crowding of 
septa, partly because of delicacy of septa and of distended gut 
wall in material often not well preserved. Even in specimens 
carefully fixed in a relaxed condition, variation in location of the 
supposed valve was found. This perhaps should have been antici- 
pated because of variation in gizzard location. The data secured 
from laboratory as well as field-preserved material were insuffi- 
cient to show correlation between location of gizzard and valve. 
Situation of the supposed valve, in desmogasters and hastirogas- 
terSj as far back as the region xxv-xlii, suggested more careful 
examination of structure throughout the entire anterior portion 
of the gut. That, unfortunately, soon became impossible. How- 
ever, if gizzards are at the beginning of the intestine, the esopha- 
gus has been extended in some lines of moniligastrid evolution 
from xii to xxv, 2 or through more segments than in bringing the 
gizzard of the supposed haplotaxid ancestor of the megadrili from 
v back into xii. There may then be no need for assuming dis- 
appearance of a more anterior gizzard and evolution of other, 
and perhaps intestinal, gizzards behind gonad segments. 

Moniligastrid vasa deferentia, though often longer than in any 
other family, open at or close to the intersegmental furrow next 
behind the funnel septum. During their growth the male ducts 

2 An imlivi'hui] series of 1MU moniligasti'iil gizzanls may be anywhere in xii-xxx. 



GATES ; BURMESE MONILTOASTRIDS 303 

pass (always?) into the segment in front of the funnel septum. 
In certain circumstances the duets conceivably could acquire ex- 
ternal openings in that metamere so as to be in front of the 
testis septum. As an individual aberration (Drawida sp.) male 
pores sometimes are closer to the testis septum and so are in the 
position characterizing series Lumbrieulina in Michaelsen's Pro- 
sopora. Situation of male pores does not now appear to be a 
proper character for defining" suborders and their sections or 
series. One rather common and widely transported lumbricid 
even has the pores in front of the female apertures in agreement 
with the moniligastrids. 

Moniligastrid prostates also are different from glands bearing 
that name in many megadriles. The structure appears to be 
more like that of the eudrilid euprostates except for presence on 
the coelomic face of a glandular tissue, Moniligastrid and eudri- 
lid prostates usually have been said to be enlarged terminal 
portions of the male deferent apparatus which is correct only 
from the viewpoint of superficial appearance. Male gonoducts 
in the Qligochaeta (so far as is known) grow away from funnel 
rudiments and eventually to or through the parietes. Prostates, 
except in the Megascoleeidae (where mesodermal origin is diag- 
nostic), probably always an 1 centripetal growths from the body 
wall. Proof of that origin, even in absence of information about 
developmental stages, is provided for the Eudrilidae as well as 
the Moniligastridae by occasional presence in adults of "pros- 
tates" that are not associated with male gonoducts, 

Spermatheeae of three moniligastrid genera, though monaxial, 
as in various microdrile and megadrile families, always are dis- 
tinguished by location of the ampulla, dorsally. Additionally, the 
duct always is longer than the distance between ampulla and 
sperniathecal pore and often is much longer, a condition for which 
no explanation has been proffered. A secondary axis, when pres- 
ent (Drawida and Moniligaster) 3 always lacks a special seminal 
chamber and so is different from the diverticulum of acanthodri- 
lid, octoehaetid and megaseolecid spermatheeae. 

Most moniligastrid peculiarities were recognized in field-pre- 
served material. As microscopic anatomy is studied in proper 
preservation, other typical characters are likely to be found, 
especially in the excretory system but perhaps also in the muscu- 
lature. Formal expression of the uniqueness and isolation from 
other earthworms in more elaborate classifications would seem to 
require, as was recognized by Yamaguchi (1953), recognition 
of a taxon Moniligastrina. 



304 BULLETIN : MKSEUM OF COMPARATIVE ZOOLOGY 

Ovisacs, always dorsal, at maturity often extend back through 
several segments and are filled (in the many specimens that 
were examined by the author) not with ova but with granules of 
several sizes. These particles, long' believed to be yolk, are of 
unknown chemical nature. After the breeding season the granules 
gradually disappear and the formerly distended sacs collapse. 
Massive production of yolk now appears to be another monili- 
gastric! specialty. 

The vascular system, in spite of some apparent generic dif- 
ferences with regard to the hearts, does conform strictly to a 
family pattern, as the last pair always is two segments in front 
of the ovarian metamere. Genital ducts, male and female, open 
to the exterior variously but always in conformity with the pat- 
tern. This means, in effect, that moniligastrid genera still are 
distinguishable, as in the past, only by location of the gonads. 
In contrast, the status of each genus similarly distinguished in 
other families now 7 is suspect. 

Rather large segment numbers now appear to be characteristic 
of the moniligastrids. The smallest desmogaster (86-115 by 4-5 
mm.} has more than two hundred metameres, and the smallest 
drawida (30-41 by 1,5 mm.) has more than a hundred. 

Habitats. Moniligastrids were believed (ef. Stephenson, 1930, 
p. 608) to need "moister conditions than any other family of 
earthworms, taken as a whole, M presumably because of lack of 
dorsal pores — "usually connected with an aquatic habitat, ?? The 
moisture requirement was deduced from casual site records and 
absence of species in the drier regions of India. Dorsal pores are 
indeed lacking in the limicoline Criodrilidae and Sparganophil- 
idae but also are absent in the Hormogastridae as w r ell as two large 
families, the Eudrilidae and Glossoscolecidae, that are mainly if 
not entirely terricolous. Almost any sort of earthworm is likely 
to be found at supposedly typical drawida sites, mud, under 
stones in water, in saturated soil by a spring, etc, during periods 
of drought. One introduced drawida now is established at Lahore 
where precipitation is less than twenty inches a year, hi the dry 
zone of central Burma drawidas are common but the war pre- 
vented publication of data as to habitats, Burmese desmogasters 
certainly are not limicolous and probably most drawidas also are 
terricolous. 

Distribution. India to Korea on the Asiatic mainland, Japan, 
Borneo, Sumatra. Absence of endemic species in the Philippines, 
Java and other Malaysian islands, even the Andamans, should 




GATES: BURMESE M0NILIGASTRID3 305 

not be assumed until earthworms have been collected more svs- 
tematically in those areas. 

Tlie range, with the exception of Sumatra, is the same as of 
the best known genus, Dmwida. Ranges of other genera are 
much smaller and except for that of the South Indian Moniligas- 
ter apparently with internal discontinuities. However, few more 
than a hundred specimens (even if as many as that) of Eupaly- 
{/aster, Desmogaster and Hastirogaster have been studied. 

The distributions provide little support for the classical belief 
that Dmwida is the youngest genus, or for its descent from 
Moniligaster. 

Genus DlUWIDA Miehaelsen 1!)00 

Sites of the testes, in the smallest juveniles that have been 
examined, were indicated by opacities in septum 9/10 near to 
or about at the usual ventral position of oligochaete gonads, i.e., 
near parietes and mesially; The septal thickening is larger in 
longer juveniles and higher up in the body. At maturity, the 
sac now markedly protuberant from the septum, usually is just 
above the gut. 

Although the male pore is only one segment behind the testis 
septum, the vas deferens always is much longer than the distance 
between gonad and aperture, occasionally a hundred times or 
more longer. The gonoduct, as it leaves the male funnel, often 
passes downward for a short distance in a schizocoelic intraseptal 
space but apparently always emerges into the coelomic cavity 
of the preseptal segment where it is farther away from the site 
of its future opening to the exterior. Within that anterior 
metamere continued growth seems, judging from adult con- 
ditions, to have been random, looping the duct back and forth, 
up and down, even around the heart. Eventually, the duct 
passes back through the gonad septum and always enters the 
prostate of its own side. Before attaining that junction in some 
species, the duct penetrates into the body wall where it may join 
the prostate or turn back into the coelomic cavity. As the duct 
always reaches its proper gland, regardless of presence of simi- 
lar glands nearby, a later portion at least of its growth seems 
no longer to have been aimless but directed. Control presumably 
is provided by some sort of attraction emanating from the pros- 
tate. If that influence is exercised while prostatic anlage are 
confined to the parietes, junction may take place there, after 
which some prostates, by their own further growth, carry an ectal 



306 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

portion of the gonoduct back into the eoelomic cavity again. 
Junction with other prostates appears to be acquired only after 
gonoduets and glands had emerged independently into the coe- 
lom. In such cases, attraction presumably still emanated from 
growing ental portions of the anlage which had gotten into the 
coeloni before gonoduets had acquired junction. Unexplained, 
however, is the apparently invariable passage of gonoduets from 
the funnel septum into the anterior segment where they are far- 
ther away from the level at which they will finally open to the 
exterior- Return of the gonoduct to the postseptal metamere, 
on the contrary, could be attributable to prostatic attraction. 
Whether the gonoduct reaches its greatest elongation in the 
eoelomic cavity of x before or after junction with the prostate 
cannot now be stated. 

The prostates, according to early accounts, have at least 
three layers, a middle muscular one, an inner glandular, and 
an outer glandular, the latter derived either from the inner layer 
or from the peritoneum. Origin of the inner glandular layer 
was not mentioned but doubtless is ectodermal. The middle layer 
of moniligastrid prostate?;, at least in early growth stages, may 
well be muscular but in adults of various drawidas is often thin, 
translucent or transparent, sometimes reddish and brittle. The 
color, as in the clitellum, may have been u developed " after 
preservation but the peculiar texture seems "unlikely to have 
resulted from a post-mortem modification of muscle tissue. 
Whether muscular or otherwise, the middle layer gives to the 
prostates a wide variety of shapes, one of which seems to be 
characteristic in each species. The outer glandular layer com- 
pletely covers the eoelomic face of the capsule in some species, 
but in others is more restricted to an ental portion or to some 
rather definite zone or area. 

Digitiform prostates, present in some species of each genus 
except Moniligaster, have been thought to be primitive. GM 
glands of one drawida, identical in structure with the supposedly 
primitive prostates except for the blind ental ending, were 
thought to show descent of the bitestieulate Drawida from the 
quadritesticulate Desmogaster. The latter genus has testes in 
septa 10/11-11/12 but extra prostates in the supposedly derived 
Drawida were so placed that testes wonld have been in septa 
8/9-9/10. More recently, prostate-like glands were found (in 
other species) in segments vii-ix as well as in x where, along- 
side the real prostates, they are supernumerary. An extra gland 



GATES : BURMESE MOJSTILIGASTRIDS 307 

i?i x, closed entally just like t Ik* glands in vii-ix, obviously can- 
not 1)0 "the thickened terminal part ,? of a male gonoduet. Such 
an appearance, however, can he achieved if gonoducts during 
their centrifugal growth penetrate into apices of developing 
capsular glands. 

The "prostates" in some species of Drawid-a (and also of 
Dcsmoffftsfcr) have no glandular investment but whether these 
" muscular ? * glands (?) arc 1 capsules that have lost the outer 
glandular layer is unknown. 

Spermatheeae of some drawidas are monaxial but ducts are 
slightly widened within the parietes. A similar thickening pre- 
sumably was the rudiment from which a secondary axis was 
developed in various lines. This still reaches, in an early stage 
of evolution, only a little above the body wall but now is joined 
on one side by the slender spermathecal duct, A more obvious 
secondary axis, as well as an ectal portion of the male termi- 
nal ia in various microdriles, long has been called an atrium. 
Spermathecal atria are thin-walled organs that may be digiti- 
form or saccular and much longer or thicker than the main axis, 
or even thick-walled chambers that sometimes become bifid. 
Each atrial bifurcation, in a species group not otherwise pres- 
ently distinguishable from Drawida, bears a dichotomously 
branched outgrowth that presumably is glandular. The atrial 
aperture of other species, in a distinctly demarcated genital 
marking, has been withdrawn into a preatrial ( !) parietal in- 
vagination. Sperm rarely have been found in the atria and then 
only in an ectal portion. Function of the spermathecal atria 
is unknown but is unlikely to be the same as that of spermathecal 
diverticula in other megadriles which have a more or less dis- 
tinctlv demarcated seminal chamber solely for storage of the 
sperm received in copulation. 

Ovaries of all drawidas always are in segment xi (abnormality 
excluded). This character, uniquely diagnostic among earth- 
worms, greatly puzzled classical oligochaetologists, and has 
further consideration in a subsequent section. 

Distribution. The natural range, as already noted, is that of 
the family, with exclusion of Sumatra, and is much larger than 
that of the supposedly all conquering megascoleeid Phereiima. 
Transportation, presumably by man, has resulted in the follow- 
ing additions to the natural range: Aru, Soemba, Caroline 
Islands, Bahamas, and Puerto Rico. 



308 BULLETIN : MUSEl/M OF COMPARATIVE ZOOLOGY 

Taxonomie Characters 

The latest key to species of Drawida (Stephenson, 1923, pp. 
127-130) has 48 couplets or triplets for 42 of the 43 species then 
known. Even so, all data except that in the original descrip- 
tion was disregarded in entering Z). glmtensis Michaelsen 1910, 
Many other species, it is now clear, would have been equally 
refractory if specimens additional to very short series or the 
unique type had been dissected. Brevity of early descriptions 
(often still un supplemented) and immaturity (usually unrecog- 
nized) of types, in which definitive genital structure was incom- 
pletely developed or even undifferentiated, added to the diffi- 
culties. 

Several classical characters, such as shape and position of 
testis sacs, shape of prostates (except when muscular), latitudi- 
nal location of female pores, presence or absence of dorsal pores, 
now are known to have no taxonomie validity; Relative widths 
of intersetal intervals, as Stephenson admitted, may differ from 
one part of the body to another as well as from one worm to 
another and also may vary aecor diner to decree of contraction 
or relaxation at preservation. Other characters are of little use 
because of limitation to presence or absence, as in the case of 
pigment and peripheral closure of ovarian chamber, or because 
condition of material often does not allow a decision as to which 
of those pairs is involved. The taxonomie value of characters 
provided by gizzards (number, location) and genital markings 
(number, location, shape, size), even after accumulation of con- 
siderable data, still remains to be determined. 

Shape of muscular prostates, latitudinal location of sperma- 
thecal and male pores, as well as atrial characters, do have 
some, more or less limited use. Spermathecal pores may be at 
B, mBC or C but in Burmese worms are, with one exception, at 
the lateral level. Male pores may be at B (one Burmese species) 
or between B and mBC where differences are too small and vari 
able to be useful. Spermathecal atria may be subject to con- 
siderable intraspecific modification. Thus, a slight symmetrical 
thickening of an ectal portion of the spermathecal duct has be- 
come, in the northern part of the caerulea range, a muscular 
diverticulum that is larger than the original axis. Great intra- 
specific variation in length of digit if onu atria has been found 
in two of the better known species. Thin -walled saccular atria, 
on the contrary, are of about the same shape and size in a 
number of species. 



GATES: BURMESE MONILIGASTRIDS 309 

The characters allowed to have taxonomic value by classical 
authorities, as Stephenson's key proves, were too few to enable 
recognition of relationships. Other characters now appear to 
be of greater significance. Length of male gonoducts may be 
distinctive in some species but confirmation is required as great 
intraspecific variation has been found in two of the better known 
forms. Entrance of male gonoducts into parietes prior to junc- 
tion with prostates, emergence into the eoelom before joining, 
intraparietal junctions that are concealed or obvious, now appear 
to be specifically invariable, but ducts of all Burmese species 
enter prostates directly, i.e., in the eoelom without first passing 
into body wall. Shape of capsule in glandular prostates seems 
to be free of intraspecific variation but in manj' Burmese species 
is digitiform. More ectal portions of the deferent apparatus 
can provide a number of characters. Male pores in an appar- 
ently primitive stage are superficial, minute and recognizable 
with difficulty due to absence of differentiated structure around 
them. Often, areas containing the pores are protrusible in more 
or less characteristic shapes. Such porophores in a retracted 
state, and especially when delimited by deep grooves, have a 
rather discoidal appearance. Differences between states of com- 
plete protrusion and retraction, when unrecognized as such, 
have been thought to distinguish species (cl. synonymy of nepah 
nisis). Adequate characterization still is impossible for some 
Burmese forms, in spite of examination of numerous specimens, 
because one or the other of the states has not been seen. Primary 
male pores may be invaginate and then the larger superficial 
apertures provide another character. Existence of parietal in- 
vaginations is suspected in certain species* In others a short 
ectal portion of the prostatic duct may prove to be eversible as 
a sort of penis. Deeper invaginations that were ignored or un- 
detected by earlier authors also furnish important characters ; 
penial bodies, tubular penes, genital markings and even glands 
may be present in chambers that reach into the eoelomic cavities. 
Determination of relationships between muscular prostates and 
the eoelomic copulatory chambers, as for instance in D. caerulea, 
may provide further characters. 

An ectal portion of the spermathecae may repay more care- 
ful examination than it usually has had in the past. The pri- 
mary spermathecal pore of one Burmese species is invaginated 
and the shallow parietal chamber contains a distinctly delimited 
genital marking, 



310 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Glands associated with epidermal genital markings provide 
a set of characters in addition to absence. Tubular glands, once 
thought to be prostates, have obvious though minute pores that 
were thought to be male pores. Size, relative to that of the real 
prostates, may be specifically distinctive. Solid glands have no 
lumen and presumably no aperture. The wall may be trans- 
parent to translucent or opaque and then is soft or tough, Shape 
and size, especially of the tough-walled glands, may vary in a 
single specimen but the condition of the outer layer now ap- 
pears to be uniform within a species. Clear glands (with trans- 
parent walls) apparently do not become as large as the others 
and may be buried so deeply in tire parietes as to be unrecog- 
nisable from the eoelom without removal of most or all of the 
longitudinal musculature. 

The excretory system, as in other supposedly holonephridial 
families, has had very little attention, Earlv ontogenetic abor- 
tion of nephridia in segment ii may prove to be common if 
not universal in the family. Later disappearance of organs in 
one or more segments of the genital region now seems likely 
to provide a set of taxonomically useful characters. Nephridia 
arc vesiculate and the bladder is a diverticulum for caecum) 
given off from the tubule shortly before entering the parietes 
in the few moniligastrids for which information has been re- 
corded. Oaeeal bladders may prove to be characteristic of the 
family. Parietal portions of nephridia! ducts now seem more 
likely to be of interest to systematists. Duets enter the body 
wall near the ventral setae of postgenital segments in several 
species of Desmogaster and H astir og aster. Entry, in anterior seg- 
ments of a Japanese drawida, now is near the lateral setae 
though still at the low r er level in postgenital segments. In two 
Burmese drawidas, for which information still is available, the 
parietes is entered, in postgonadal segments, at tlie lateral level. 
Within the body wall ducts may or may not pass directly to an 
epidermal aperture. In the latter case ducts grow ventrally or 
dorsally for some distance before finally turning towards the 
epidermis. Irregular alternation of nephropores between two 
or even three levels, usually with more or less asymmetry, has 
been found in several species. Nephropore locations, accordingly, 
need not indicate levels at which ducts enter the body wall. 

Number of segments in types usually lias been recorded. One 
type, 47 by 1.75 mm., has 150 segments. Another, 55 by 2 mm,, 
has 165. The only record of less than a hundred segments (98) 




GATES : BURMESE MONILIGASTRIDS 311 

is of a unique specimen that may bo a posterior amputee- One 
Indian species was said to have more than 500 segments and 
five others have more than 300 or 400. Information available 
as to segments of n r pal c n sis (129-180), gracilis (140-340) and 
other species does not suggest hatching with a definitive number. 
Much tedious counting doubtless will be required to determine 
specific limitations. 

Color characterizations of the past often are of little or no 
importance to the taxonomist today. An unpigmented lumbrieid 
species, Octolasium cyaneum (Savigny, 1826), by its name pro- 
vides one bit of supporting evidence. Another luinbrieid exam- 
ple is furnished by Eisensia rosea (Savigny, 1826) which usu- 
ally appears to be unpigmented though minute yellowish or 
brownish (epidermal?) flecks often are recognizable under the 
binocular in older individuals. Colors sometimes mentioned, or 
occasionally shown in plates, are attributable to cuticular iri- 
descence, blood, even to ingest a within the intestine or (espe- 
cially at posterior end of the body) to accumulations of coelomic 
corpuscles and debris. A green color, apparently characteristic 
of a few species, cannot be traced to discrete particles. A beauti- 
ful red color of the clitellum in many drawidas is "developed" 
after preservation by formalin and perhaps also by other sub- 
stances. The fine granules responsible for that color are in the 
outermost portion of the epidermal cells. Similarly located 
granules may he responsible for a striking orange or red colora- 
tion of the clitellum in live specimens of E. rosea but after 
preservation the pigment flecks appear to be yellowish or 
brownish. 

Pigment usually is in or is associated with the circular muscle 
laver of the bodv wall but may extend into the longitudinal 
layer at the anterior end and dorsal] y. Presence of granules 
that appear to be red or reddish brown in sections through the 
body wall does not always confer a similar color. Such worms 
often seem to be blue, sometimes even a quite dark blue or 
almost black. The clitellar region, at maturity, of pigmented 
drawidas after preservation may be white though pigment is 
present underneath the epidermis. 

A clitellum rarely has been recognized in vivo and monili- 
gastrids were for a time placed in a group called Aclitelliens. 
The tumescence that has sometimes been noted in preserved 
specimens may have been developed by the preservatives. Ex- 
cept as interrupted by genital markings, the clitellum probably 



312 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

is always annular in the Moniligastridae. Though intergeneric 
differences have been recognized they do not now appear to be 
of taxonomic importance. 

Certain characters not mentioned in the taxonomic section 
as they now appear to be universal throughout the family are : 
absence of typhlosoles, supra-intestinal and calciferous glands; 
location of extra-esophageal blood vessels lateral to the hearts; 
presence of a subneural trunk that is adherent to the parietes 
rather than to the nerve cord as in the Lumbricidae* Seminal 
vesicles , possibly present in some ancestral form before direc- 
tion of proliferation by the testes was reversed, always are 
absent. Penial and copnlatory setae never have been found. 

Drawida beddardi (Rosa) 

1890. Moniligaster beddardii Rosa, Ann. Mus, Sto. Nat. Genova, 29:379. 
(Type locality, Chiala, Burma. No types.) 

1894. Moniligaster beddardi^ Bourne, Quart. Jour. Micros* Sei*, 36:374. 

1895. Moniligaster barwelli (part), Beddard, A Monograph of the Order 

of Oligoehaeta, p. 200. 
1900. Drawida barwelli (part), Michaelsen, Das Tierreich, 10:116, 

1923. Drawida barwelli (part), Stephenson, (The Fauna of British India), 

Oligochaeta, p. 133, 

1924, Drawida banvelli var. hehoen&is -\- D. *"* fluvai tilis ** Stephenson, Rcc. 

Indian BIus.j 26:324, 325. (Type localities, TIeho plain and White 
Crow Stream, both near Ynun^whe. Types in the Indian Mus,) 

1926. Drawida tecta Gates, Idem, 28:148, (Type locality, Yanngwhe. 
Paratype in XL S, Natl, Mus.) 

1931. Drawida hehocnsis f Gates, Idem, 33:340, 

1933, Drawida hejionixis, Gates, Idem f 35:443. 

The type locality of this species was said to be "Villagio di 
Chiala 14004500 m (Carin Padaung o Asciuii Ghecu)." The 
district mentioned was not a political division and reference 
presumably was to some area inhabited by Padaungs seventy- 
odd years ago when Fea was collecting in Burma. Tt may have 
been almost anywhere between Leiktho Circle and Karenni. The 
village probably passed out of existence long ago. No informa- 
tion about it was obtainable in Burma and a better guess than 
that above could not be made bv Italian missionaries stationed 
at the area in question. 

All drawidas found on the Shan Plateau during 1925-1940 
were referred to three species, kehoensis, longatria and nepal- 
ensis. The latter two are clearly distinguished from beddardi 



GATES: BURMESE 1I0NILIGASTKIDS 313 

by length of their male deferent ducts and by the genital mark- 
ings as well as associated glands. The color mentioned by Rosa 
provides no clue and may have been unnatural (preservation 
was alcoholic). The dorsal pores supposedly present doubtless 
were the "small dark spots" noted by Stephenson in his ma- 
terial. The "labbra rigonfie" of the type may well have been 
tumescences such as occasionally were recognizable in hchoensis, 
around the secondary male apertures. With these clarifications 
there remains no morphological or distributional contra-indica- 
tion to suppression of hehoensis as a synonym of beddardi. 

D, barwelli, with which beddardi was mistakenly synony- 
mized, was never found in Burma. 

Each male pore of the classical authors opens into an in- 
vagination comparable to copulatory chambers of various phere- 
timas. The much smaller primary pore is at the free end of a 
penis pendent from chamber roof. The penis is protrusible 
through the secondary aperture and the chamber itself is ever- 
sible. Male porophores at maximal protrusion, presumably as 
during copulation, have the penes projecting from a thicker 
basal portion. External apertures of the spermathecae, no longer 
minute, are large enough to admit a distal portion (at least) 
of the porophore into what appears to be a thickening of a 
parietal portion of the spermathecal duct. This chamber has 
been called an atrium but the structure bearing that name in 
many species of Drawida, although much larger, does not serve 



as a vagina 



Drawtda bullata Gates 1933 

1933. DrawitJa fucosa Gates, Rei\ Indian ilus., 35:439. (Type locality, 

Kalewa, Burma, No types.) 
ProfflBj September, 2-26-4, K. John. 
Laboo (Prorne), September, 00-13. K. John. 
Thanbula. (Thayetmyo), September, 12-30 46* K. John. 
Taungdwingyi (Magwe), August, 0-0-12. K. John. 
Minbu, August, 13-27-10. K. John. 
Ye-U (Shwebo), August, 13-27-10. Saw San Thwe. 

External characteristics. Size (juveniles excluded), 65-95 by 
4-5 mm. (Minbu), to 94 by 4 mm, (Taungdwingyi), 55-65 by 
3-4 mm. (Ye-U), 84-120 by 4-5 mm. (Laboo), 90-180 by 4-7 
mm. (Thanbula), 150-180 by 4-7 mm. (Prome). Unpigmented, 
Nephropores, present from iii, on anterior margins of segments, 
at or close to D. Clitellar coloration ., red, extending into ix 
and xiv. 




314 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Spermathecal pores, very small transverse slits, just median 
to 0. A marginal area, forming a circumferential lip, is clearly 
marked off peripherally and often is so protuberant that the 
pore appears to be located on a transversely elliptical, small 
papilla. This usually seems to be segmental as it is bounded 
posteriorly by 7/8 but may be situated exactly on 7/8 with no 
more indication of belonging to vii than to viii. Onlv an an- 
terior portion of such a lip is marked off, on most Minbu worms, 
by a transverse groove, concave posteriorly, that does not pass 
into 7/8. These lips or protuberances may be only slight ever- 
sions of the spermathecal ducts rather than definite porophores. 
Female pores, minute, circular, in transverse areas of greyish 
translucence, at or just lateral to B. 

Male pores, transverse slits on ventral ends of whitened, 
anteroposteriorly flattened, rather pointed protuberances in BC f 
with median margins nearer to B than lateral margins are to C 
and often reaching B. Each porophore is bounded posteriorly 
fay the presetal secondary furrow of xi and in front apparently 
(most specimens) by 10/11 which curves anteriorly around the 
porophore. On other worms 10/11 seemingly is continued just 
onto the median and lateral faces of the porophore where it 
becomes unrecognizable, the porophore bounded anteriorly by 
a groove that opens into 10/11 just beyond those blind endings, 
On such worms the male pore is in line with 10/11 and the 
porophore accordingly appears to be formed from parts of both 
x and xi though that from x is much the smaller. Porophores 
on other worms appear to belong wholly to xi, the male pores 
much nearer anterior than posterior margins. The porophore 
is delimited anteriorly, in Taungdwingyi worms, by a trans- 
verse furrow near posterior margin of x that does not pass into 
10/11 but here also the larger part of the porophore obviously 
is from xi. Solid or other definite glands always are lacking in 
porophores of this species. 

Genital markings, whitened areas of epidermal thickening, 
distinctly delimited as a rule but not associated with special 
glands, in (viii)-ix-xii (cf. Table 1), Markings of x usually 
are paired : transversely oval and with pointed end mesially, 
in median half of BC but sometimes reach A or into A A, in 
the postsetal annulus (Minbu, Taungdwingyi) ; longitudinal, 
reaching to or almost to B and C and through set al -postsetal 
annuli (Prome, Laboo, Thanbula, Ye-TJ) or confined to post- 
setal annulus (1, Ye-U). Paired markings of xi are transverse, 



GATES; BURMESE MONILIGASTRIDS 315 

usually smaller than in x, in a median part of BC but occasion- 
ally reaching A, confined to seta] or to setal-postsctal annuli. 
Unpaired and median markings are transverse, usually extend- 
ing through setal-postsetal annuli (in which case the presetal 
annulus is quite short ) t reaching well into BC or even to C. 
A pair of markings that clo not quite meet at mV, in viii of 
one Laboo worm and in ix of one Minbu worm, if united would 
be of the same size as the unpaired markings. On the right 
side of a Thanbula worm the marking of x is small and trans- 
verse but the marking of xi is large and longitudinal, the re- 
verse of the usual condition as well as of that on the left side. 

Internal anatomy. Gizzards, 1-3, in xv-xviii (Table 2). Three 
gizzards of a Prome specimen that may prove to belong to this 
species are in xviii-xx. 

Testis sacs, usually un constricted, about equally in ix and x, 
Vas deferens, slender, usually iridescent in 9/10 and in ix, in 
a number of short loops on anterior face of 9/10 median or 
close to or around hearts of ix. Occasionally, further loops 
(Minbu) are bound by connective tissue in a small cluster 
against posterior face of 9/10. A more eetal, non-iridescent 
and seemingly thickened portion in x, less than 5 mm, long, 
is slightly sinuous, zigzagged or in several short loops and 
passes into ental end of prostate directly. The cluster of hair- 
pin loops of the thickened portion in x of worms from other 
localities may be about half the size of the testis sac above it 
(Ye-U, Prome), smaller to nearly as large (Thanbula), as large 
to larger (Laboo, Taungdwingyi). The thickened portion in a 
Ye-IT worm is 60+ mm. long. Prostates, protuberant into coelom- 
ie cavity and usually bent over towards the nerve cord, narrowed 
only within the parietes, ental end bluntly rounded, short- 
ly elliptical in cross section, occasionally with an appearance 
of slight anteroposterior flattening, glandular investment con- 
tinued to body wall. Capsule, usually reddish, soft, digitiform, 
slightly curved, 1 (Taungdwingyi, Minbu) to 2 mm, (Prome, 
Thanbula) long, 

Spermatheeal atria, digitiform, in vii. Atria are erect on 
anterior face of 7/8, irregularly constricted or very shortly zig- 
zag-looped and less than 5 mm. long (Minbu), or are in a clump 
of shortly u-shaped loops bound to ventral parietes. The cluster 
may be small and ventral or large enough to reach to or almost 
to dorsal parietes. Length, 10-20 mm. (Ye-U), 10-12 (Prome), 
15-20 (Taungdwingyi) j 60-70 mm. (Laboo), A terminal por- 
tion about 10 mm. long in some Laboo worms is filled with 




BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

a sticky white material in which slight spermatozoal iridescence 
is visible. 

Ovarian chamber, closed off from parietes. Ovisacs, extend 
into xiv or xv. 

Juveniles. Greyish translucent spots, anlago of male pores, 
are at 10/11 in the two smallest juveniles on which no poro- 
phores or protuberances are recognizable. Male porophores of 
larger juveniles are represented by slight swellings of anterior 
margin of xi without definite posterior demarcation, the pores 
now apparently behind 10/11 which seems to be bent forward 
slightly around the protuberances. If segment x contributes any 
part to the male porophore it certainly must be much smaller 
than that from xl 

Juveniles still without genital markings have male gonoducts 
of about the same length as those of acl Stellate and clitellate 
worms from the same locality. 

Remarks. The smallest adults are from a southern portion 
of the species range where there is considerable rainfall. The 
largest w r ere found in regions of much less precipitation. 

Some aelitellate worms (3 Taungdwingyi, 2 Ye-U, several 
Thanbula) appear otherwise to be fully mature* 

From the postseptal portion of male gonoducts, in worms 
softened after long formalin preservation, a sheath had sepa- 
rated off so as to reveal internally a slightly zigzagged or sinu- 
ous tube of about the same thickness as the preseptal portion. 

Male porophores have been seen only in a protuberant state, 
presumably just as at copulation. Information as to the com- 
pletely retracted state is needed, 

D. fucosa was distinguished from hullata by quantitative 
differences such as soma size, length of spermatheeai atria and 
of male gonoducts, Individuals as small (20-45 by 1*4-2 mm.) 
as most of those on which hullata was erected are unlikely to 
copulate with worms as large as the types of fucosa (130-170 
by ti-7 mm.) but intermediate soma sizes herein reported make 
retenlion of fucosa unnecessary. Variation in lengths of sper- 
matkeca] atria and vasa deferentia parallels that recorded below 
Tor longatri® from which hullata is distinguished by absence of 
solid GM glands. Relationships to vulyans remain uncertain. 

Abnormality, (No. 1.) Left spermatheca, with two ampullae 
{distended by white material), one attached to posterior face of 
7/8 in usual manner, other hanging down in coelomic cavity, 
each with a discrete duet V/ 2 mm. long, the united duct passing 
vent rail v as usual 




GATES: BURMESE MONILIGASTRIDS 



317 



Table 1 



Frequency distribution 


of 


seg 


omental 


position of 


median 


and 


paired 


genital 


ma: 


rkings in D. 


b allata 














Median 


t 




Paired 














Segments 












* 


X 


xi 


xil 


X 


xi 


Prome (30) 














30 


30 


Laboo (13) 






7 




3 


l 


13 


10 


Thanbula (76) 






11 




4 


19 


76 


72 


Taungdwingyi (12) 






12 








12 


12 


Minbu (37) 






25 






8 


37 


37 


Ye-U (9) 






3 


2 




3 


7 


9 



Figures in parentheses, number of specimens from the locality. 



Table 2 

Frequency distribution of segmental position and number of 

gizzards in D. bullata 







TABLE 2 




Locality 


XV 


Segments 
xvi xv M 


xvii 


Taungdwingyi 


5 


12 




Ye-U 


9 


11 2 




Minbu 


14 


27 28 


2 


Laboo 


2 


11 12 


3 


Thanbula 


2 


15 18 


7 


Prome 


5 


17 18 


12 



X umber 

1 


of tfiz/.unls 
2 3 


7 


5 




11 




13 5 




8 4 




15 4 




5 14 



Drawida caerulea Gates 1926 

Kayan (Hanthawaddy), August, 0-5-8. K. John. 

Thongwa (Hanthawaddy), August, 0-6-8. K. John. 

Thinbawgyin (Bassein), October, 0-0-6, K. John. 

Pegu, August, 0-0-2. ** Jungles to the east/' 0-3-1, K. John. 

ThanatpLn (Pegu), August, 0-2-3. K. John. 

Toungoo, October, 0-2-0. K> John. 

Minbu, August, 0-0-2, K„ John. 

Myingyan, September, 0-0-2. K, John. 

Myotha (Sagaing), September, 0-1-1, K. John. 

Ava (Sagaing), September, 0-3-6. K. John, 

Maiictalay, September, 0-7-0, K, John. 

Ye-U (Shwebo), September, 0-9-34, Saw San Thwe. 

External char act eristics. Nephro pores, present from iii. Left 
male porophore of one Kayan worm is completely retracted 
though the right is completely everted. Genital markings, al- 
ways lacking. 



:{18 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Internal anatomy. Pigment, always present in the body wall. 
Gizzards, one or two only in Ye-U specimens, with one excep- 
tion and then three in xiv-xvi. Commissures from extra-eso- 
phagcals, behind 8/9 (60 specimens). 

Vas deferens, slender, iridescent, twisted into a small cluster 
of loops that are firmly bound together (southern localities), 
slightly thicker throughout but still iridescent and loops much 
less firmly bound together (Ava), slightly thickened and iri- 
descent throughout but longer and loops easily separated, length 
65-70 mm. (Ye-U). 

The sperm a thecal duet of southern worms obviously is more 
thickened in a terminal portion than in species of the beddardl 
group. The thickened part is pyriform or almost conical and 
mainly in the body wall. In Ye-U worms the thickened portion 
is asymmetrical and reaches forward halfway to 6/7 and presum- 
ably must be called an atrium. Size of the atrium is less than 
half that of rasUis in which 6/7 and dorsal parietes are reached. 

Ovisacs may extend as far back as into region of xix-xxii. 

Abnormality. (No. 2,) Left male porophore, at 9/10. Left 
male organs, one segment anterior to usual location. Ovarian 
chamber, in two discrete halves, on the left side one segment 
anterior to usual location as is the female pore. An ovisac ex- 
tends posteriorly from each ovarian chamber, Left spermatheca, 
lacking. 

The left mesoblastie somite at the eighth level presumably 
was aborted during early embryonic development and its place 
was taken by the one at the ninth level. 

Regeneration. Tail regenerate, of eight segments at 136/137. 

Remarks. Data as to number and location of gizzards in 
specimens listed above were lost, as well as records that would 
have permitted characterization of prostates more adequately 
than in the past. 

Male porophore* of all previous southern specimens were 
fully protruded. Maximal protrusion and complete retraction 
now has been shown by a single worm (Kayan). The thick- 
walled muscular chamber into which the terminal half of the 
protruded porophore is withdrawn has been considered to be 
a prostate. This chamber is distinguished from the prostates 
of most moniligastrids by absence of a glandular layer on its 
coelomic face. Pending acquisition of further information about 
both kinds, the term prostate with characterization of *' muscu- 
lar M is retained. 




GATES : BURMESE MONILIGASTRIDS 319 

Whether a porophore as large as that of southern worms can 
be inserted^ without change of form, into the spermatheeal atrium 
seems doubtful. The ejaeulatory apparatus certainly appears to 
be highly evolved even though a discrete penis (such as is pres- 
ent in heddardi) is lacking. The atrium in rasilis is much larger 
than is required for reception of the porophore of southern 
worms but the protruded condition has not been seen in northern 
worms. Atria of decourcyi Stephenson, 1914, are even larger 
but here again only the retracted state of the male porophores 
has been seen. 

Atrium now seems a more appropriate characterization for the 
parietal invagination into which the spermatheca opens through 
an aperture on a discoidal genital marking. The primary sper- 
matheeal pore may prove to be within the atrium and if so the 
external aperture will be tertiary rather than secondary. 

Southern worms were distinguished from rasilis mainly by 
quantitative differences such as length of vasa deferentia and 
size of spermatheeal atria. The range of D. caerulea caerulea 
as now known is from the Irrawaddy delta to latitude of Manda- 
lay. D. caerulea rasilis was found in the C hind win valley near 
Monywa. Worms from the region of Mandalay to Ye-U and 
Monywa are intermediate with respect to one or another of the 
characters distinguishing the two subspecies. Considerable 
searching through difficult territory much farther north may be 
required to reveal whether intermediate forms exist between 
caerulea and decourcyi. 

Drawida delicata nom. no v. 

19H0. Dmivida sp„ Gates, Bee, Indian Mus. f 32, 298, 

Remarks. A filament at distal end of each male porophore 
may be only cuticular lining of the spermiducal passageway 
that had been loosened and then evaginated during strong con- 
traction at preservation. Whether such contractions protruded 
discoidal porophores normally extending into x-xi or everted 
small parietal invaginations is unknown. A "blueish shade" 
may not have been associated with pigment any more than in 
the unpigmented lumbrieid, Octolasium eyaneum (Savigny, 
1826). The "minute" size of spermatheeal ampullae and short- 
ness of ovisacs (in xii) may or may not be indicative of im- 
maturity. 



320 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

The species is distinguished from beddardi by the greater 
length of male deferent ducts, by absence of parietal invagina- 
tions at ectal ends of male deferent apparatus or, if male poro- 
phores are everted chambers, by absence therein of protrusible 
tubular penes* Elongation of the male gonoducts also distin- 
guishes delicata from all other Burmese species with adiverticu- 
late spermathecae except spissata. The latter requires no 
consideration because of great differences in spermathecae and 
prostates. Relationships presumably are with species yet to be 
collected in the little known areas west and south of Mergui 
district, 

Drawtda flexa Gates 1929 

Zinlia (Tavoy), September, 0-0-1. W.D.Sutton. 

Siyigyan (Tavoy) f "plains'* September, 0-1-3. W. D. Sutton. 

Kawletelmung (Tavoy), "nearby hills," September, 0-4-5. W. D, Sutton. 

Pyinthadaw (Tavoy), "nearby hills, M September, 0-2-1. W. D. Sutton. 

Nyaungdon (Tavoy), September, 0-0-1. W. IX Sutton. 

Migyaunglaung (Tavoy), "plains," September, 0-1-0. W # D. Sutton. 

Thaton, September, 0-0-2. K. John. 

Duyinzeik (Thaton), September, 0-14-7, K. John, 

Xaunggala (Thaton), September, 0-3-3. K, John, 

Bilin (Thaton), September, 0-15-12. K. John, 

Taungzung (Thaton), September, 0-8-15, K. John. 

Kinmunsakhan (Thaton), September, 0-0-7. K, John. 

Boyagyi (Thaton), September, 0-1-0. K. John. 

Kyaikto (Thaton), September, 0-12-4. "Nearhy hills/ 1 September, 0-8-15. 

K. John. 
Sittang (Thaton), October, 0-2-0. K, John. 

External characteristics. Size may be rather small, as in 
worms from Kyaikto hills, 50-80 by 3-4 mm. (diameter measured 
across clitellum which is much wider than the rest of the worm). 
Nephropores, present from iii, in anterior margins of segments, 
usually at or close to D, pores of viii always dorsal to D, pores of 
ix usually at or only slightly dorsal to D but occasionally one or 
both dorsal and, more rarely, even farther from D than in viii. 
Dark spots at mD and near some preclitellar intersegmental 
furrows may look more or less like dorsal pores but epidermis 
and cuticle are imperforate even when there seems to be a gap 
in the musculature. Clitellar coloration, lacking vent rally on x 
and a presetal portion or all of xi, from a level just beyond 
lateral margin of male pore lips. Laterally from that level the 
epidermis is quite tumescent. 




GATES: BURMESE MONILIGASTR IDS 321 

Spermatheeal and male pores, larger than uephropores, cer- 
tainly larger than female pores and accordingly not minute, 
margins of pores almost never smooth. Spermatheeal pores, just 
median to 0, at 7/8 or (apparently) just in front of 7/8, Male 
pores, at 10/1 1 ? close to mBG, longitudinally or transversely 
slit-like or irregular, each at center of an indistinctly delimited 
but nearly circular, white and slightly tumescent area reaching 
to or nearly to B 9 to a tertiary furrow on postsetal secondary 
annul us of x and the preset al secondary furrow of xL The poro- 
phore is usually divided into two portions by 10/11 but when 
tumescent the furrow may be indistinct or even unrecognizable, 
especially close to the pores. The portion in front of 10/11 is 
lacking in Sittang worms. 

Genital markings, usually transverse, 2 to 20. located as fol- 
lows; Median (unpaired and rarely longitudinal), presetal in 
viii and x-xii, postsetal in vii-x, occasionally in setal annul i of 
vii and x. Paired, in EG, presetal in viii-xii, postsetal in vii and x, 
in setal annul i of viii-xi, in median half of BC (postsetal mark- 
ings of vii, presetal of x-xi ? those in setal annulus of x), or with 
centers nearer mBC, Duyinzeik and Taungzun worms have 
16-20 markings each. 

Internal anatomy. Gizzards, three (4 specimens), four (55), 
five (10), in xvi-xxiii (Table 4), Commissures from cxtra- 
esophageals, in front of 8/9. 

Vas deferens, short, slender, and for most part iridescent 
(clitellate specimens), in several loose loops on anterior face 
of 9/10, twisted around heart of ix, several further loops on 
posterior face of 9/10, an ectalmost portion 1*2 mm. long ap- 
parently slightly thickened and certainly without recognizable 
iridescence. Prostates, erect, recumbent or held against parietes 
by delicate strands, J- or U-shaped, entalmost portion of capsule 
only slightly or not at all widened. 

Spermatheeal atria ? small, usually 1-2 mm. long, in Taungzun 
worms occasionally reaching a length of 3 mm. An ental por- 
tion, about half the length or slightly less, usually distended 
and with thin wall. Lumen gradually narrowed and wall thick- 
ened in ectalmost portion. Spermatheeal duct, passing into pos- 
terior face of atrium in vii close to parietes. 

Reproduction. Ampullary coagulum of clitellate worms usu- 
ally is characterized by a rather brilliant iridescence presumably 
spermatozoa!. Spots of iridescence also are clearly visible in 
the coagulum within distended ental portions of the spermatheeal 
atria in several Taungzun worms, Iridescence of male gonoduets 



322 



BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 



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GATES : BURMESE MONILIGASTRIDS 



323 



presumably is due to presence of sperm on the way out from 
the testis sacs. As sperm are matured and exchanged in copula- 
tion, reproduction is assumed to he sexual and bi parental. 

Regeneration. The only record now available is of a tail re- 
generate, at 170/171, with terminal anus and 36+ segments. 

The difference in thickness of ectal and ental portions of 
spermatheeal atria, and of the shape, is so little that the organ 
can be called digitiform. A similar slight distention of an ental 
portion sometimes was recognizable in much longer atria of 
bullata and longatria. 

The flexa range in Burma appears to be restricted to the Tenas- 
serim division and there from Tavoy north to the Sittang River. 
The Thailand boundary may not be a natural boundary for the 
species. 

Relationships are with longatria from which flexa is dis- 
tinguished by the smaller genital markings, smaller size of the 
glands associated with those markings, absence of a gland in male 
porophores and shortness of spermatheeal atria. Atria are of 
about the same length in several races of longatria but not in 
those found in the flexa range. Additonal differences may be 
recognizable when retracted states of longatria male porophores 
can be seen. Although differences are quantitative, all specimens 
from the Tenasserim division have been referable, without ques- 
tion, to one or the other of the species involved, 



Table 4 

Frequency distribution of segmental position and number 

gizzards in D* flexa 



of 



Locality 


xvi 


x\"ii 


WWW 

XV] 11 


Kpgi 
xix 


n puts 

XX 


xxi 


xxif 


Numbe 
xxiii 3 


r of 
4 


gizzard? 
5 


Duyinzeik (17) 


i 


6 


1 *"> 

1 _ 


17 


17 


15 


3 




2 


10 


5 


Taungzuii (22) 




6 


17 


21 


22 


18 


6 




1 


18 


3 


Pyinthadaw (3) 




2 


3 


3 


3 


1 








3 




Naunggala (6) 




1 





5 


6 


6 


_ 




1 


3 


2 


Kawletehaung (9) 




4 


8 


8 


9 


5 


1 


i 




9 




Sivlgyan (4) 




1 


4 


4 


4 


3 








4 




Miscellaneous* ( 8 ) 




2 


7 


8 


8 


6 


1 






8 




Totals 


i 


22 


56 


0(5 


09 


54 


13 


i 


4 


55 


10 



* Includes : xvii-xx (1 Mig.vaunglau.ng, 1 Thaton), 

xviii-xxi (2 Sittang, 1 Thaton, 1 Boyagyi, 1 Zinba), 
xix-xxii (1 Nyaungdon). 




324 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Drawida gracilis Gates 1925 

That on, September, 23-0-0. K. John. 

Kyaikto (Thaton), September, 1-0-0, K. John. 

Thongwa (Hanthawaddy), August, 24-4-0. K. John. 

Twante (Hanthawaddy), September, 1-0-0, K. John, 

Hlawga (Insein), September, 4-0-0, K, John, 

Wanetchaung (Insein), September, 5-0 0. K. John, 

Taukkyan (Insein), September, 7-1-0, K. John. 

Thinhawgyin (Bassein), October, 0-3-0, K.John. 

Pegu, August, 38-0-0. "Jungle to the east/ 1 60-0-0. "Jungle to the west," 

33-0-0, K. John. 
Thanatpin (Pegu), August, 9-0-0, K, John, 
Paukkanng (Prome), September, 9-0-0, K. John. 
Prome, September, 11-0-1. K. John. 
Thanbula (Thayetmyo), September, 5-3-0, K, John. 
Sandoway, "Hills," September, 19-0-0. I. M. Ismailjee. 
Ramree (Kyaukpyu), "nearby hills," September, 1-0-0. I. M. Ismail jee. 

External characteristics. Nephropores, present from iii, Clitel- 
lar coloration, red, lacking ventrally on x and presetal half of 
xi. Male porophores of the clitellate specimen are protuberant, 
rather conical but with rounded distal ends, firm but containing 
no glands. Porophores of less mature worms are more nearly 
discoidal but possibly because adult organization had not been 
completely achieved. 

Genital markings, indistinctly delimited areas of epidermal 
thickening, paired, transverse, between B and a level just beyond 
lateral margin of male porophores, in the presetal annulus of 
x, anterior portion of the postsetal annulus of x, setal annulus 
of xi but extending forward into presetal annulus and occa- 
sionally almost, to the intersegmental furrow. Markings in x 
of the clitellate specimen meet mesially, a slight furrow along 
mV the only indication of a boundary. The ventral setae are in- 
cluded in the anterior markings. The epidermis in a presetal 
portion of ix (Prome, clitellate) or viii and ix (Thongwa, aclitel- 
lates) is thickened in BB or even CC but no markings are recog- 
nizable externally, A longitudinal marking, in a median portion 
of BC (3 Pegu specimens), extends through the whole length of 
x as on one of the types. 

Internal anatomy. Gizzards, two (1 specimen), three (26), 
four (6) ? in xiv-xx (Table 6). Intestinal origin, in xxi (speci- 
men with gizzards in xiv-xvii), just behind 21/22 ( specimens 
with gizzards in xiv-xvii and xiv-xvi). Commissures from extra- 
esophageals, in front of 8/9, 



GATES: BURMESE MONILIGASTRIDS 325 

Vas deferens, slender, rather long, twisted into two clusters of 
closely compiled loops, one on earh face of 9/10, clusters of 
about the same size or the anterior slightly larger, the two to- 
gether smaller than the testis sac, slender throughout. Prostates, 
sessile on parietes, of circular outline, capsule small, 0.5-1.25 
mm, long, shortly ovoidal, pointed end within the parietes. 

Sperinatheeal atria, digitiform, 1.0-1.5 mm. lonpr, lumen nar- 
row, an irregular cleft in transverse section. Ovisacs (clitellate 
worm) reaching into xvii. 

Ah nor nut! if}/. (No. 2.) Male porophores, two pairs, on 10/11 
and 11/12, each porophore with a minute, transversely placed 
male pore. Greyish translucent spots in 11/12 at B just median 
to margins of male porophores are about at sites of female pores. 
Testis saes of 9/10 are normal but in the region where 10/11 
should be there is on each side of the body a fairly large cluster 
of loops of a second vas deferens that is not connected at all with 
the anterior male gonoduct. The prostate, on each side, is a 
single acinous mass extending slightly across levels of inter- 
segmental furrows 10/11 and 11/12. The anterior vas of a side 
passes into the anterior margin, the second into the posterior 
margin of the prostate. Paired ovisacs extend into xiv. 

Ovaries and oviducal funnels were not distinguishable, in 
part perhaps because of poor preservation. Relationships of sep- 
tum 10/11 to 11/12, to the gut ? and to the parietes were not 
determined. 

Gonads of xi may have been hermaphroditic. 

Remarks, Length of fully mature worms is unknown, since 
only anterior portions of aclitellate and clitellate individuals 
were obtained. Absence of mature specimens in collections from 

Table 5 

Frequency distribution of segmental position of genital 

markings in D, gracilis 

Segments 
Locality x xi 

presetal post seta 1 

Thongwa 3 8 8 

Taukkyan 6 6 

Prome 7 7 7 

Thanbula 2 2 2 

IJlawga 2 2 2 

Pegu 14 4 

Paukkaung 3 3 3 

Thinhawgyin 3 3 



326 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

so many localities may have been due to failure to dig deep 
enough. 

Some geographical variation in a range as large as that of 
gracilis would not he unexpected. 

Table 6 

Frequency distribution of segmental position and number of 

gizzards in IX gracilis 



Locality 


xiv 


XV 


Spfrnie-nts 
xvi xvii xvlii 


xix 


XX 


Number 

o 


of pizza rrls 
3 4 


Thanbula (3) 


3 


3 


3 










3 




Thongwa (7) 


3 


7 


7 4 








1 


5 


1 


Pegu ( 15 ) 


2 


6 


11 14 


10 


5 


2 




10 


5 


Prome (5) 




4 


5 5 


1 








5 




Taukkyan (1) 






1 1 


1 








1 




Thmb&wgyin (2) 






1 2 


2 


1 






o 




Totals 


8 


20 


28 26 


14 


6 


— 


1 


26 


6 



Dhawida longatrta Gates 1925 

External characteristics. Nephropores, obvious, present from 
in, except on x of adults, at or very close to CD except in viii 
where they are somewhat more dorsal, A longitudinal dark band 
of rather blueish appearance (as if pigment were showing 
through the epidermis) usually is visible at niD except in the 
most anterior segments. Also visible at inD, close to various 
anterior intersegment al furrows, are markings that look more 
or less like dorsal pores, 

Internal anatomy. A discrete longitudinal muscle band at 
mD is lacking. Instead, the longitudinal muscle layer gradually 
becomes thinner toward niD and just at thai level is almost non- 
existent. This thinning is responsible for the externally recog- 
nizable dark band at mD. Close to intersegmental levels at mD, 
especially anteriorly, after stripping off the longitudinal muscu- 
lature, there are visible slight protuberances or depressions but 
cuticle and epidermis are not perforated, ie, ? dorsal pores are 
lacking. 

The gut behind the pharynx is rather slender, in viii-x with 
low but lamelliform longitudinal ridges on inner wall. The gut 
is valvular, in individuals of the nominate race having gizzards 
in xv-xviii, through all of xxii or of xxiii. 

The dorsal trunk is single anteriorly until disappearance into 
tissues of the pharyngeal bulb. The subneural, usually large 



GATES : BURMESE MONILIGASTKIDS 327 

and blood-filled, is adherent to the parietes. Even before re- 
moval of the cord, the trunk usually is recognizable in contracted 
specimens as closed ends of the loops protrude 1 beyond each side 
of the cord. Nephridia, apparently vesiculate behind the clitol- 
lum, bladders perhaps elongately sausage-shaped (condition poor 
and relationships of parts not determinable with certainty). 
Nephridial ducts, behind the clitellum at least, pass into parietes 
at CD gap. Nephridia were not found in x of adults and pre- 
sumably are aborted prior to maturity. Brain and commissures 
to sub pharyngeal ganglion were left posteriorly in iii by a trans- 
verse section exactly along 3/4, 

Glands associated with genital markings have no lumen but 
are provided with a thick, tough and obviously muscular wall. 

Remarks. As male porophores always have been more or less 
markedly protuberant the retracted states cannot, yet be charac- 
terized. Because of appearances in specimens with less protu- 
berance it is suspected that the primary male pores may be 
mvaginated slightly in a state of complete retraction. 

Supposed pores of GM glands, previously recorded, are now 
believed to have been artifacts. These *' glands/' without any 
lumen and thus called solid, may have mechanical rather than 
secretory functions, such as stiffening penes (certain Chinese 
Species) or male porophores {longatria) , gripping: penes or poro- 
phores during copulation when present in walls of spermathecal 
pore invaginations (some Chinese species) or in the parietes close 
to the spermathecal pores (longatria) , 

Internal anatomy of all specimens available since 1941 is like 
that of the nominate race as characterized above* 

Variation in shape, size and direction of male porophores, in 
size, shape and location of genital markings, is so great that 
attempts to define a few externally identifiable varieties w r ere 
abandoned long ago. Many large collections from the recent 
alluvium in the deltas region south of Henzada-Tharraw r addy, 
secured after 1932, contained only individuals that were like 
the Rangoon types on which the species was erected. This ma- 
terial enabled recognition of a location pattern for genital mark- 
ings though many individuals lack one or more of the set. An 
area somewhat larger than that just indicated had been de- 
limited, before Pearl Harbor, as the probable range of a nominate 
race. Presence of the same form in places as far apart as Ran- 
goon, Burma and Palembang, Sumatra, must be attributed to 
overseas transportation, presumably by man. The original source 



328 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

obviously is Burma, Since habits are such as to permit transpor- 
tation, and colonizing ability had been demonstrated, it was then 
possible to assume that presence of the nominate race in isolated 
Burmese localities such as Myitkyina, Bhamo, etc., likewise is a 
result of transportation. Disregarding such colonies which ap- 
peared to be localized in and immediately around towns to which 
potted plants are known to have been taken from Rangoon, de- 
limitation of ranges of other races, usually found in less urban 
situations, was under way when the Japanese invasion ended 
local study of Burmese earthworms. 

The races differ from one another in various characters, some 
of which are determinable only from dissection. Some of the 
differences have been thought to distinguish species, Male poro- 
phores probably are not exactly the same in any two races of 
which there are an unusual number. 

Drawida lostgatria long atria Gates 1931 

External characteristics. Segments, 183 and 206, the last few 
mctameres of each worm very small, without externally recog- 
nizable setae but showing no evidence of regeneration, 

Spermathecal pores, not minute, transversely slit-like* 

Genital markings: unpaired, median, presetal, in AA, of xi 
(6 specimens), xii (16), xiii (7) ; paired, each in median half 
of BC in xii (3), united marginally with a median marking (2), 
or rudimentary (1). An area of greyish translucence, in lateral 
half of BD or even reaching beyond D, in preset al annulns of 
viii lacks the distinctly demarcated rim of the other markings 
but is associated with a characteristic parietal gland. A less ob- 
vious area of epidermal modification, associated with a small 
gland, is present in the posterior part of vii just median to each 
spermatheeal pore. 

Regeneration. Three of the sixteen worms from Bhamo and 
Myitkyina, with 124, 171 and 172 segments, are unregenerate 
posterior amputees. Another individual had lost its hind end 
at time of collection. Ten of the twelve remaining specimens 
have tail regenerates. Number of segments in the regenerates; 
at 104/105, 86 setigerous + several very short and without ex- 
ternally recognizable setae; at 124/125, 23 + 7 or more; at 
136/137, 14 + 6 or more ; at 143/144, 16 + 6 or more ; at 165/166, 
6 + ?, The anus in each of those cases is terminal. Young re- 
generates at 142/143, 145/146, 151/152 and 152/153 each have 
a dorsoterminal anus that may reach nearly to the substrate. 



GATES: BURMESE M0N1LIGASTK1DS 329 

Segments, usually without, externally recognizable setae, usually 
are distinguishable only on the ventral side. The distance from 
substrate to dorsal margin of the anus, in a regenerate with 
indications of 10 + segments, is only about a quarter of the 
distance from substrate to ventral margin. One regenerate, at 
196/197, is unsegmented. 

Remarks. The two unamputated worms, with 183 and 206 
segments, obviously had become sexually mature before meta- 
meric differentiation had been completed in a terminal portion 
of the body. 

External characteristics. Length of worms from Tharrawaddy 
district, to 170 mm . Diameter, to 7 mm. Segments, 149 4- a 
number of rudimentary metanieres without externally reeogniz- 
able setae, 177 + 1 or 2, 190 + several (3 specimens), 191 + 2 
or more, 193 + 1 or % 206 + 2 or 3, 210 + 2, 211 +2 (2 speci- 
mens) s 220 + several. 

Genital markings: unpaired, median, presetal, in AA of xii 
(2 specimens) ; paired, each in median half of BC or reaching 
to A, presetal, in xii (32), in viii where margin is indistinct as 
in Bhamo-Myitkyina worms (41), occasionally replaced by two 
or even three smaller markings, postsetal, in lateral part of BC 
in vii (41) and there small, even more indistinct than in viii. 

Regeneration* Three worms are unregenerate posterior am- 
putees, Thirteen worms had lost their hind ends at or since 
time of collection. Six of the remaining twenty-five have tail 
regenerates as follows: still unsegmented, anus reaching for- 
ward in dorsum nearly to substrate; anus dorsoteriniual, 
segmentation indicated ventrally but setae unrecognizable exter- 
nally ; at 134/135, 10 setigerous segments + a number on which 
setae are unrecognizable, anus dorsoterminal : at 143/144, 12 + 
several; at 191/192, 3 + several; at 197/198, 5 + several 

Remarks. Each of the 19 unamputated worms has, imme- 
diately in front of the anal segment, one or more circumferential 
furrows that presumably are intersegmental. In the axial por- 
tions thus delimited, nephropores and setae are unrecognizable. 
The portion of the body in front of the anus then is a growth 
region in which metameric differentiation had not been com- 
pleted W'hen the worm became sexually mature. 

These Tharrawaddy worms probably were secured in the 
plains. 



330 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Drawida longateia verrucosa Gates 1931 

External characteristics. Segments, 205, 206, 213, 224, 225 ? 
233, 237, 238, in each case with two or more rudimentary seg- 
ments already demarcated but without externally recognizable 

setae, 

Spermatheeal pores, like the male apertures, small transverse 
slits, Male pores, facing anteriorly or anterolateral^. Tumes- 
cent margin of the male pore with a slight but definite groove 
coming out of the pore and passing posteriorly on ventral face 
of the tumescence. Male porophores, in AD, reaching equators 
of x-xi or shortened at either end. Each disc-like porophore may 
have an anterior and a posterior genital marking, or markings 
may scarcely be distinguishable though glands are present, or 
either marking and the gland may be absent. 

Genital markings, additional to those of the male porophores, 
are as follows. Unpaired, median, in A A, presetal in viii (3 
specimens), in setal annulus of viii (3). postsetal in x (11), xi 
(13), xii (17). Paired, presetal in AA of x (2, in AB of viii (3), 
x (13), xi (2), postsetal in x (15) ; larger and in some part of 
AD on eaeh side, postsetal in vii (17), postsetal in viii (15, in 
two of which each marking is replaced by two smaller ones), 
presetal in x (4), presetal in xii (14). 

Regeneration. One worm is an unregenerate posterior ampu- 
tee and another had lost part of its tail at time of collection. 
Four have tail regenerates. One has no externally recognizable 
segmentation. Another, also with a dorsoterminal anus has sev- 
eral rudimentary segments (without setae) marked off ventrally. 
Two regenerates, at 198/199, have each a terminal anus; seg- 
ments, 6 ( + ?) and 19 (+?). 

Remarks. Each of the eleven unamputated worms has sev- 
eral rudimentary segments at its posterior end. 

This series of seventeen specimens from Tharrawaddy district 
probably was secured in the hills or in jungles remote from 
the town. 

Drawida nana Gates 1933 

The GM glands have a soft, opaque wall without muscular 
sheen and may represent an intermediate stage in evolution 
from the "clear" sort with transparent wall to the lont/afria 
sort with a strongly muscularized wall. If, however, definite 
pores are present in the associated genital markings, as was origi- 
nally believed, the glands will not be solid and will provide 
additional evidence for distinction from longatria. 




GATES ; BURMESE MONILIGASTEIDS 331 

Drawida nepalensis Michaelsen 

1907, Drawida nepalensis Miehaelsen, Mitt. Nat ur hist, Mus. Hamburg, 
24:146. (Type locality, Gowehar, near Katmandu, Nepal. Type 
in the Indian Mus,) 

1909* Drawida nepalensis + D. burchardi Miehaelsen, Mem. Indian Mus., 
1:147, 149. 

1916. Drawida jalpaigiirensis Stephenson, Rec. Indian Mus., 12:307. (Type 

locality, Jalpaiguri, at base of the eastern Himalayas* Type in the 
Indian Museum.) 

1917, Drawida nepalensis, Stephenson, Rec. Indian Mus., 13:372. 

1922. Drawida 'nepalensis , Stephenson, Idem, 24 : 430, 

1923. Drawida nepalensis + D. burchardi + B* jalpaigurensis, Stephen- 

son, (The Fauna of British India), Oligochaeta, p. 146, 134, 141. 

1924. Drawida troglodytes Stephenson, Rec. Indian Mus., 26:129, (Type 

locality, Siju Cave, Garo Hills, Assam, Type, in the Indian Mus,) 

1925. Drawida bur char di + D. hodgarti + D. papillifer (part), Stephen- 

son, Idem, 27:50, 51. 

1926. Drawida eacharensis Stephenson, Idem, 28:251. (Type locality, Kat- 

licherra, South Oachar, Assam. Types, in the Indian Mus.) 

1929. Drawida nepalemis, Stephenson, Idem, 31 : 229, 

1930. Drawida nepalensis, Gates, Idem, 32; 290. 

1931. Drawida nepalensis, Gates, Idem, 33:348. 

1933. Drawida burehardi, Gates, Idem, 35:426. 

1934. Drawida nepalensis + D m troglodytes, Gates, Idem, 36:242 and 253. 

Andaman Islands 

Port Blair, Station E 10, 2 specimens. Station B 15, 0-1-0, (Indian Mus.) 
Station 6, 1-0-0. (Indian Mus.) 

(No station indicated), September 1932-May 1933, 0-1-0. H. S. Rao, No 
data, 0-1-0, (Indian Mus.) 

Burma 

Sandoway, riverside, September, 0-1-0. Hills, September, 0-6-1. I. M. 

Ismail jee. 

Akyab, September, 0-16-17. I. M, Ismailjee. 

Myohaung (Akyab), September, 0-0-7. I. M. Ismailjee. 

Naba (Katha), nearby hills, September! 0-2-0, Saw San Thwe. 

Lashio (Northern Shan States), 7th mile on the Namtu road, September, 
0-3-15. Wan Hu Mone village, 5 miles from Lashio, muddy ground 
covered with water cress, September, 0-0-1, II. Young. 

E Nai village (North Hsenwi State), 9 miles from Lashio, at base of bam- 
boo clumps, September, 0-5-20, H. Young, 

Bhamo, September, 0-4-8. K. John* 

Myitkyina, September, 0-0-14, K. John. 

Weshi (Myitkyina), October, 0-3-0. F, D, Forbes. 




332 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

India 

Lokia ( Balipara Frontier Tract, Assam), S.xi.1939, 0-2-0, S. L. Hora (In- 
dian Mus.) 
Teesta Bridge, Teesta Valley, 2/6/34, -40-1. 8, L. Hora (Indian Mua.) 
Nagrota (Kulu District, Punjab), garden of P. W. D. rest house, 13/6/26, 
0-1-0. S* L. Hora (Indian If us.) 

Pakistan 
Lahore and vicinity, 29*16-13. H. K, Bhalti. 

External char act eristics. Size (Pakistan), 78-105 by 4-5 mm. 
Segments (Pakistan), 86 (posterior amputee), 113 (p. a*), 114 
(p. a.), 129, 130, 137, 138, 139, 140, 144, 153, all juvenile, 135, 
145, 148, 150, 154, 156, 157, 158, 159, 161, 162, 163, 164 (2 speci- 
mens), 166 (2), 168, 169. 173, 176, aclitellate and clitellate speci- 
mens. Setae (Pakistan), closely paired, AB = CD, A A > BO, 
DD ca. — ^C, unrecognizable in last three or four segments. 
Nephropores, present from iii, at D except slightly more dorsal 
on vii (Pakistan) or vii-viii (Burma), usually lacking (or un- 
recognizable?) in x ( Burma) or x and xii (Pakistan) though 
quite obvious in xi. A pore definitely is present on right side 
of xii (1, Pakistan) and vestiges of pores of x sometimes become 
visible after treatment with picric acid. 

Male porophores of most specimens are as previously described 
by the author and in a condition that must now be regarded 
as of maximal or near maximal protrusion, presumably as dur- 
ing copulation. Porophores of one Pakistan worm, almost if not 
fully retracted, are circular to shortly elliptical areas distinctly 
delimited from x and xi, depressed slightly below general epider- 
mal level. Segments x and xi are slightly indented so that short 
anterior and posterior portions of the porophore are concealed 
from view. More indentation, if further change is possible, pre- 
sumably would result in complete coverage of porophores which 
might then appear to be in some sort of a chamber. 

Genital markings are of at least two sorts. 1) Small, circular 
areas of greyish transl licence, one in each male porophore and 
one in vii just in front of each spermathecal pore. Around each 
of the anterior markings there often is a fairly wide and opaque 
band of slight epidermal thickening. The translucent area is 
the outer face of a nearly spheroidal solid "gland" filled with 
greyish translucent tissue. Porophore markings occasionally are 
not recognizaMe though the glands are present. Anterior glands 
may bulge the body wall up into the coelom or may be visible 
internally through gaps in the musculature. Glands of vii were 




GATES : BURMESE MONILIGASTRIDS 333 

not found in several Arakan specimens but epidermis at sites of 
markings is slightly tumescent and wrinkled. 2) Areas of slight 
epidermal modification and not associated with solid or other 
glands. Two small transverse markings (Burmese worms) in each 
setal annulus of x and xi, about in line with the male porophores, 
epidermis slightly thinned, A median, transversely elliptical 
marking (many Pakistan worms) reaching A or B on each side 
in presetal half of xi, epidermis slightly thickened and without 
elitellar coloration. Paired, translucent areas of epidermal thick- 
ening in lateral part of BO or reaching D t presetal in vii 

( Lashio ) . 

Five Arakan worms have a median presetal marking of vari- 
able shape and size in AA of ix. A central portion of each is 
translucent. Removal of the longitudinal musculature disclosed 
in the remaining thin portion of the body wall only a translucent 
spot which may be a vestige or rudiment of a gland. 

Internal anatomy. A special longitudinal muscle band is 
lacking at mD where there is a gap in the musculature from 3/4 
or 4/5 posteriorly that is of about the same appearance as those 
at setal levels. The gap is slightly widened just behind inter- 
segmental levels. Removal of the longitudinal musculature dis- 
closes a slight protuberance at mD just behind level of each 
intersegmental furrow. The protuberance is over site of a some- 
what pore-like marking which is behind rather than at inter- 
segmental level. The cuticle and epidermis are imperforate, 
hence there are no dorsal pores, 

A low rather broad median ridge is present on the floor of 
the esophagus in middle segments of the region behind the last 
gizzard. Gizzards, two to four (Table 7), in types of cacharensis 
are located as follows: xiv, xvi, xviii (2 specimens) ; xiv-xv, xviii 
(1) ; xiv, xvii-xviii (1) ; xiv, xviii-xix (1) ; xv, xviii (1) ; xvii, 
xix (1)* Intestinal origin (Lokra worms), in xxiv (1 specimen), 
xxv (1), xxvi (3), xxvii (5). Enterosegmental organs in four 
or five postgizzard segments are especially obvious. 

Dorsal blood vessel, probably complete but traceable only into 
iv. Ventral trunk, complete, bifurcating over subpharyngeal 
ganglion, the branches traceable along the circumpharyngeal 
nervous commissures nearly to the brain. Subneural trunk, ad- 
herent to parietes, traceable anteriorly only into x or ix. Com- 
missures from extra-esophageals, in front of 8/9. From each 
posterior commissure a vessel may pass back on dorsolateral 
aspect of gut into xii or xiii, with four transverse connections 



334 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

to the correspond in g vessel of the opposite side. One of those 
paired vessels usually is unrecognizable, perhaps because it is 
empty rather than absent. The vessel that is visible is near the 
median plane and presumably is the one believed in the past to 
be a supra-esophageal. Hearts, of vi-ix lateral, Nephridia, lack- 
ing in x, vesiculate, bladder (usually called a caecum) elongately 
sausage-shaped. Nephridia! duets, pass into parietes at D, 

One testis sac occasionally is displaced posteriorly underneath 
the ovarian chamber (13 Akyab specimens, neither sac displaced 
in 22 Akyab worms), Vas deferens, slender and iridescent in 
0/10 and ix. with several small loops on anterior face of 9/10, 
one loop encircling heart of ix, thickened in x and there opaque, 
twisted into a cluster of loops that may be as large as the testis 
sac, Prostatic capsule^ slenderly club-shaped, only very slightly 
and gradually widened entally, 2-4 mm, long. 

Spermathecal atria, 3-5 mm, long, reaching into contact above 
gut or overlapping slightly, an ectal portion of variable length 
stalk-like, the widened ental portion usually irregularly con- 
stricted, Spermathecal ampullae (Pakistan), empty, translu- 
cent. Ovisacs (Pakistan), apparently reaching back into xiv-xv 
but actually within pockets of 12/13, Contents of distended 
ovisacs consist of yolk granules of several sizes. Ova, if at all 
present, certainly must be very few (aclitellate as well as clitel- 
late worms). Female funnels, vertically elongated and band-like, 
on posterior wall of the chamber from parietes to opening into 
ovisac, lateral margins folded over toward each other so as to 
form a sort of trough almost closed. Ovaries (preservation poor) 
apparently band-like and placed vertically opposite the female 
funnels. No egg strings were found. 

Juveniles, Male porophores, on most of the Teesta juveniles, 
are represented by very small swellings on which the pores but 
not the genital markings are visible. Bach swelling is demarcated 
anteriorly as well as posteriorly by a slight transverse furrow 
that, does not pass at either end into 10/11. The latter is con- 
tinued, on each side of the porophore, nearly to the tip and the 
male pore. Deepening of intrasegmental grooves and extension 
to 10/11 along with obliteration of 10/11 in the porophore pre- 
sumably would produce the appearance characteristic of adults, 
of belonging neither to x nor xi. The male pores now are assumed 
to belong morphologically at 10/11, with the porophores originat- 
ing about equally from x and xi. 

Testis sacs, in the smallest Pakistan juveniles, just below the 
gut, gonoduet loops about at sit* 1 where the testis would be in 




GATES: BURMESE M03STILIGASTR IDS 335 

Other families. Ovisacs in juveniles of some size still are un- 
recognizable and the ovarian chamber apparently is not closed 
off peripherally. Oviducal funnels are recognizable but ovaries 
were not found. 

Abnormality. Five helieoinetameres are present in the intes- 
tinal region of a Pakistan worm. 

Regeneration, Head regenerate of six segments at 6/7 
(Myitkyina). Tail regenerate, 5 mm. long (Arakan). 

Parasites. Crescent-shaped, uninucleate protozoans are pres- 
ent (Arakan worms) in numbers on the dorsal face of the gut 
near the dorsal blood vessel. 

Remarks. Variation in segment number is similar to that of 
longatria. 

Septa from 10/11 posteriorly though membranous are strong 
enough to allow considerable anteroposterior movement of a 
post genital portion of the gut — in one Pakistan worm the gut 
of xii-xiv and including a gizzard is in front of the ovarian 
chamber. Preservation, in the case just cited, was such as to 
permit tracing septa to insertions on gut as well as parietes thus 
enabling recognition of morphological location of the gizzard. 
Septa in the postgenital region often are adherent to the gut 
beyond the real insertions which may not always have been iden- 
tified correctly. Even if all discontinuities in the gizzard series 
are mistaken the condition responsible for the erroneous deter- 
minations is characteristic of many individuals of nepalensis. 

Male porophores of adult worms previously identified as 
nepatensis by the author were markedly protuberant presumably 
as during copulation. One of the Pakistan worms now has shown 
the retracted condition of the porophores. 

D. racharensis is known only from the type series comprising 
16 (3 previously dissected) aclitellate (?) specimens* Differ- 
ences from ittpalensis are restricted to the male porophores. 
When protuberant, the porophores are like those of the author's 
nepalensis or slightly more conical. The genital marking some- 
times is unrecognizable but the solid gland is present. Retracted 
porophores are as in one of the Pakistan worms. Similar poro- 
phores of the unique type of troglodytes then are also in a re- 
tracted state. Tlier e now is no eoiitra-indieation to synonymiza- 
tion of racharensis and troglodytes. D. jalpaigurensis has been 
known hitherto only from the original account of a single aclitel- 
late specimen i4 in a bad state of preservation." Examination of 
the type some years ago provided the data recorded below* 



336 BULLETIN : Ml'SEl'M OF COMPARATIVE ZOOLOGY 

Addendum 

External characteristics. Spermathecal pores, transverse slits 
at 7/8, slightly median to C. Male pores, not recognizable. Male 
porophores, transverse, very slightly raised, reaching mesially 
near to B and laterally at least to mBC, 10/11 continued slightly 
into the median margins but not into the lateral margins. An- 
teriorly and posteriorly each porophore is bounded by a definite, 
erescentic furrow concave towards the porophore, neither furrow 
passing into 10/11 mesially or laterally. A distinctly demarcated 
circular area just behind level of 10/11 in each porophore is the 
outer face of a tough-walled ovoidal gland protuberant into 
coelomic cavity, narrower and buried in the parietes. Genital 
markings, paired, presetal in vii, postsetal in vii and there just 
in front of each spermathecal pore. 

Internal anatomy. Vas deferens, short. Glandular investment 
of the prostate, much thicker than the slender capsule. Sperma- 
thecal duct, in vii passing into posterior face near parietes of 
a saccular erect atrium. The gland of the postsetal genital 
marking of vii (labelled atrium in Stephenson 1923, fig. 51, p. 
141) protrudes slightly into the coelom. 

Remarks. Except for apparent shortness of the male gono- 
duets no evidence was found to justify retention of jalpaiguren- 

sis. 

Forma abscisa Gates 1931 

Kutkai (Northern Shan States), Dak bungalow grounds, ca. 4500 i'oi-t, 
November, 1920, 0-0-1. H. S. Rao (Indian Mus.). 

Namkhaiu (Northern Shan States), streams and pools on the north bank of 
the Shweli River, ca. 2500 feet, December 1026, 0-0-8. H. S, Rao (In- 
dian Mus.). 

Lashio (Northern Shan States), 0-3-0. H. Young. 

Myitkyina and vicinity, September, 0-4-2. K. John. 

Kadranyang (73rd mile on road from Myitkyina to Putao), October, 0-3-0. 

F. I). Forbes. 
Kawa pang (82nd mile on same road), October, 0-5-0. F. D. Forbes. 
Xawnghkai (250th mile on same road), October, 0-15-0. F. D. Forbes. 
1 1 ting bai (92nd mile on same road), October, 0-7-0. F. D. Forbes. 

External characteristics. Nephropores, present from iii. Sper- 
mathecal pores, transverse slits with smooth margins (aclitellate 
specimens) exactly on 7/8. Male pores, very small transverse 
slits but quite obvious and in concave depressions of ventral 
faces of protuberant porophores. 



GATES: BURMESE ; MONILIGASTRIDS 337 

Genital markings here also are of two sorts; 1) Associated 
with a solid gland as in ne pale u sis. Very small, circular, greyish 
translucent areas in vii-viii close to 7/8 and near the spermuthe- 
eal pores. One marking always is present in the vicinity of each 
sperma thecal pore but there may bo one or two more. 2) Circu- 
lar, opaque and indistinctly delimited markings, unpaired even 
though not median, with a small translucent central spot and 
not associated with glands. These markings are lacking on ten 
of the worms. 

Internal anatomy. Gizzards, three to five in xiii-xix (Table 9). 
Commissures from extra-esophageals, in front of 8/9. 

Reproduction. Spermathecal ampulla of previous specimens, 
even though elitellate, were in a juvenile condition. No sperm 
were found in the spermathecal atria or spermathecal ducts and 
no externally adhesive spermatophores (as in all other species 
of Drawida) ever were seen, Parthenogenesis was suspected but 
in Drawida recognition of male sterility is not as easy as in some 
other genera. Testis sac coagulum was examined for evidence of 
sperm but the data no longer are available. 

Spermathecal ampullae of six aelitellate worms from Hting 
bai, Kadranyang and Kawa pang, are large enough to be con- 
sidered of adult size. Each adult ampulla, except in one worm, 
is filled with a sticky white material w T hich may have (2 speci- 
mens) a slight iridescence that presumably is due to presence 
of sperm, However, spermathecal atria appear to be juvenile, 
ovarian chambers are empty, and ovisacs are juvenile. 

Remarks. Male porophores of all worms referred to abscisa 
were protuberant, presumably as in copulation. Though easily 
distinguishable from those of nepalensis in the protruded con- 
dition, little difference in shape or appearance is expected in 
completely retracted states. Differences of abscisa from nepalen- 
sis are mostly quantitative such as smaller size, fewer segments 
( ?), absence of a solid gland in the male porophores. The seem- 
ing discontinuitv of the gizzard series that characterizes a 
majority of specimens of nepalensis has not been found in in- 
dividuals with abscisa male porophores. Both forms have been 
transported by man and the original home of each is unknown. 
The Myitkyina records do seem to suggest that the northern 
part of the district is near if not within the original range. 
However, other earthworms collected by Mr. Forbes between 
Myitkyina and Putao arc all of peregrine species and all were 
secured in vicinity of government rest houses to which exotic 
plants in pots or tins are known to have been taken. Worms with 
abscisa characters have not been found outside of Burma. 



BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 



The locality list might well have been much longer except for 
an unfortunate rivalry. The collections from a northern part 
of the Myitkyina district were made by a missionary after the 
end of the rains when earthworm activity (cf. Gates. 1960b) is 
mostly restricted to still moist sites. Certain reptiles being col- 
lected for a museum, at the same time and in the same area, had 
to be fed on earthworms. The missionary was getting too large 
a share of the limited supply wherever he went. A complaint 
lodged with the government that the missionary was a communist 
spy resulted in an order that he return at once to Rangoon. 



Table 7 

Location of gizzards and of intestinal origin in 

Drawida nepalensis 



Specimens 



Li xiv 


XV 


xvi 


Segmi 

1 * 

xvn 


?nts 
xviii 


xix 


XX 




Locality 


1 1 






1 










Pakistan 


1 


1 




1 










Lokra 


1 


1 




1 








xxvi 


Akyab 


3 


3 




3 










Pakistan 


1 


1 






1 






xxvi 


Akyab 


1 


1 








1 


1 




Pakistan 


2 




2 




2 








Pakistan 


1 




1 






1 






Pakistan 




1 


1 


1 








xxvi 


Akyab 




1 


1 


1 








xxvii 


Akyab 




1 


1 


1 








xxviii 


Akyab 




1 


1 


1 










Lokra 




1 


1 




1 






xxvi 


Akyab 




1 


1 




1 






xxv ii 


Akyab 




1 


1 






1 


1 


xxvi 


Akyah 




1 




1 










Pakistan 




1 






1 


1 






Pakistan 






1 


1 


1 






xxvi 


Akyab 






1 


1 






1 


xxvi 


Akyab 






1 


1 






1 


xxvii 


Akyab 






1 




1 




1 


xxvi 


Akyab 






1 




1 




1 


xxvii 


Akyab 








1 


1 


1 




xxvii 


Akyab 








1 


1 




1 




Akyab 








1 




1 






Pakistan 








3 




3 


K* 


xxvii 


Akyab 


2 10 


16 


15 


20 


11 


9 


10 







gates: burmese moniligastrids 339 

Table 8 
Location of genital markings in Z>. nepalensis f. abscisa 

I i i * J I 1 1 i I p y 

Segment Intra segmental position v rp * t-,i k w I 

vi Postsetal, centered at B 1 

vii Setal annulus, in BC 1 

Postsetal, lateral half of A A 1 

Postsetal, in BC 12 

viii Presetal, in BC 1 

Setal annul us, lateral half of A A 1 

Postsetal, lateral half of AA 1 

Postsetal, centered at B 1 

Postsetal, in BC 3 

ix Presetal, centered at B 2 

Presetal, in BC 4 2 

Presetal, lateral half of A A 1 

Preaetal, median 1 

Setal annulus, lateral half of AA 1 

Postsetal, lateral half of AA 2 

x Presetal, lateral half of A A 1 

Setal annulus, lateral half of AA 1 

Postsetal, median 1 1 

xi Preset;! 1, centered at B 11 

Presetal, in BC 1 

Setal annulus, in BC 2 

Localities: N = Nawnghkai, T = Hting bai, Kd = Kadranyang, Kw = Kawa 
pa Ji£, I j = Lashio. 
Existence of usual two secondary furrows is assumed even if furrows are 
unrecognizable. 



Table 9 

Frequency distribution of segmental position and number of 

gizzards in IX nepalensis f. abscisa 



Locality 


xiii 


xiv 


sv xvi 


ts 

* ■ 

XV 11 


xviii 


xix 


Number of gizzards 
3 4 5 


Lashio (2) 


2 


2 


2 2 


2 






2 


Tingpai (7) 


1 


2 


7 7 


6 


5 


2 


13 3 


Kadranyang (2) 


1 


2 


2 2 


2 


1 




2 


Xawangkai (14) 




12 


14 14 


14 


7 




9 5 


Kawapang (5) 




1 


5 5 


5 


2 




2 3 



Totals 4 19 30 30 30 15 2 3 17 10 



340 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Drawida papillifer Stephenson 1917 
Drawida papilliper papillifer 

Bamree (Kyaukpyu), nearby hills, September, 0-5-0. I. M. Ismailjee. 

Akyab, September, 0-5-0. I. M. Isniailjee. 

Kyauktaw (Akyab), September, 51 juvenile and aclitellate specimens. I- M. 
Ismailjee. 

Butliidaung-Mauiigrtaw (Akyab), September, 0-4-0. I, M. Ismailjee. 

Paletwa (Arakan Hill Tract), September, 29 juvenile ami aclitellate speci- 
mens, I. M, Ismailjee. 

External characteristics. Nephropores, present from iii and in 
x, slightly dorsal to D in iii-vii and somewhat more so in viii, 
occasionally also in ix. 

Spermathecal pores, transverse slits, exactly on or rarely just 
anterior to 7/8, slightly median to G. Male pores, longitudinal 
or diagonal, in x though but slightly in front of 10/11, at or 
just lateral to /?, each usually in a semicircular, indistinctly 
delimited, whitened area with base at 10/11 ; occasionally a short 
transverse furrow just in front of the male pore provides an 
anterior boundary. Female pores, at B* just behind 11/12. in 
verv slight transversely slit-like crevices. 

Genital markings, small areas of greyish transhicence, circular 
or shortly elliptical (and then transverse), with very slightly 
raised and narrow, opaque rims. Markings are located as fol- 
lows: One, on posterior margin of x immediately lateral to the 
male pore (5 specimens from Akyab, 16 from Kyauktaw, 9 from 
Paletwa). As before, but just in front a second marking (20 
Kyauktaw, 10 Paletwa). A third marking on posterior margin 
of x and just lateral to the last of the other two (1, Paletwa). 
One marking just anterior to male pore (11, Paletwa), One 
marking in setal annul us of x just lateral to J3 (3 Kyauktaw, 1 
Paletwa). One marking on anterior margin of xi, on each side, 
centered in AB or at B (5 Akyab), A presetal median marking 
on xii (1 Paletwa). Other markings are transversely elliptical. 
One on vii just in front of each spermathecal pore (-5 Akyab, 
48 Kyauktaw, 22 Paletwa). One in viii just behind each sperma- 
thecal pore (4 Akyab, 46 Kyauktaw, 13 Paletwa). One in AB 
of vii, just behind equator on each side (5 Akyab). Presetal, 
on each side of vii in median part of BC (20 Kyauktaw) or cen- 
tered at mBC (11 Kyauktaw), or in lateral half of BC (12 
Kyauktaw). One in median half of BC in setal annulus of vii 
on each side (4 Kyauktaw). One postsetal on each side of vii 
in median half of BC (5 Paletwa) or in lateral half of BC (1 



GATES : BURMESE MONILIGASTRIDS 341 

Paletwa) or even dorsal to D (13 Paletwa), One in setal annulus 
of vii, on each side, dorsal to D (2 Paletwa), 

Internal anatomy. Pigment, present in circular musculature 
of dorsum even when a bluish color is unrecognizable or almost 

so externally. 

Low longitudinal or vertical ridges are present on inner wall 
of a postgizzard portion of the esophagus, Gizzards and intes- 
tinal origin, respectively, are located as follows: xiii-xv and 
xxii fl specimen), xiv-xvi and xxiii (10), xiv-xvii and xxiii (2) 
or just behnd 23/24 (7), xv-xvi and just behind 23/24 or in 
xxiv (2), xv-xvii and xxiv (3), xv-xviii and xxiv (2), xvi-xvii 
and xxiv (3). Commissures from extra-csophageals, anterior to 
8/9. From one of the posterior commissures (behind 9/10) a 
Vessel usually passes back into xii or xiii along the dorsolateral 
aspect of the gut. This vessel often is close to the median plane 
and when distended with blood probably has been mistaken in 
the past, for a supra -esophageal trunk. A corresponding vessel 
on the opposite side, in the present worms, may have been empty 
and hence unrecognizable, Nephridia are present in x. 

One testis sac is displaced posteriorly into xii or even farther 
back in 59 specimens (juvenile, sexual and postsexual). Both 
sacs are displaced in 1 sexual specimen. Neither is displaced in 
9 juveniles, 13 sexual and post sexual specimens. Vas deferens, 
short, probably about five and certainly no more than ten mm. 
long. The ental portion runs ventrally within or bound to 9/10, 
passes in front of heart of ix and then back into x where its short 
loops are in a small cluster against posterior face of 9/10 well 
above parietes. Prostates, erect, close to 9/10 so that the vas 
does not drop to or even very near the parietes though entering 
the gland below its ental end. Prostates, club-shaped, narrowed 
eetally, 2-3 mm. long (coelomi, portion), capsule rather slender 
ectally, an entalmost. portion of varying' length slightly widened 
and almost spheroidal, shortly ovoidal, or rarely almost ellip- 
soidal. 

Spermathecal atria, in vii, 2-3 mm. long. An ectal portion 
of variable length, rather slender and duct-like, may be fairly 
sharply marked off from the widened ental portion. The latter 
which may be of about the same length as, shorter or longer than 
the duet is slightly flattened and shortly or elongately elliptical 
to oval in outline. Whitish, sticky material within the ental 
portion of an atrium can be disentangled into a long cord. A 
slight iridescence (spermatozoal?) characterizes cords of one 



342 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

worm. The consilium in the spermathecal ampulla is not cord- 
like and though soft is teased apart only with some effort. 
Spermathecal duct, slightly widened just prior to passing through 
7/8, length and thickness of widened portion slightly variable. 
Segment xi always open in pinned out worms (82) and the 
ovarian chamber appears not to be closed off peripherally. 

Abnormality. (No. 2) Male pore and associated genital mark- 
ings of left side lacking. Left spermathecal pore, at 8/9. Left 
testis sac, vas deferens and prostate, lacking (Paletwa). 

Remarks. Distinguished from peguana mainly by the male 
pore locations, 

Ramree worms and three from Kyauktaw are slightly different 
from other west Burma specimens and are brief]}' characterized 
below. 

External characteristics. Spermathecal pores, apparently just 
behind 7/8, Male pores, nearer mBC than B. Female pores, 
slightly lateral to B. 

A transversely elliptical (to almost circular) genital marking 
just median to each male pore, centered at or slightly lateral 
to B and reaching A, is larger than other markings of x in these 
and all other specimens. Small translucent spots, just anterior 
and posterior or posterolateral to each male pore may be rudi- 
mentary markings. Transversely elliptical markings located as 
follows: one just in front of each spermathecal pore (3 Kyauk- 
taw, 4 Ramree) ; two in front of each spermathecal pore in a 
longitudinal scries (2 Kyauktaw) ; one just behind each sper- 
mathecal pore (4 Ramree), 

Internal anatomy. Gizzards in xv-xvii (3 Kyauktaw). Intes- 
tinal origin in xxiii (3). 

Testis sacs, not displaced. Vas deferens probably shorter than 
in other worms as the cluster of loops on the posterior face of 9/10 
definitely is smaller. Prostatic duct (portion of capsule without 
glandular investment), slightly longer, 

Coagulum in spermathecal atria, with a slight iridescence, ap- 
parently not in a long coiled cord. Atrial stalk, much shorter than 
the ental sac, 

GM gland median to male, enlarged and interrupting longi- 
tudinal musculature. 

Draw Ida papillifer peguana Gates 1925 

Migyamiglaung (Tavoy ), plains, N^pti'inlier, 0-5-0. W, I). Sutton, 
Pyinthadaw (Tavoy), hills, September, 0^0*1. W. D. Sutton, 



GATES: BFRMESE MONILIGASTKIDS 343 

Kawlet chining (Tavoy), nearby hills, September, 0-1-1. W. D. Sutton. 

Siyigyan (Tavoy), September, 0-0-6. W. D, Sutton. 

Kyaikmaraw (Amherst), August, 0-0-1. K. John. 

Thaton, Sept ember, 0-26-8. K. John. 

Duyinzeik (Thaton), September, 0-14-8, K. John, 

Xa ungual a (Thaton), September, 0-38-15. K. John, 

Bilin (Thaton), September, 0-5-6. K t John. 

Aungsein (Thaton), September, 0-6-2, K. John. 

Sit tang (Thaton), October, 0-3-0, K. John, 

Boyagyi (Thaton), September, 0-7-0, K, John. 

Kyaikto (Thaton), September, 0-12-5, K, John. Nearby hills, 0-9 8, K. 

John. 
Kyaiktiyo (Thaton), September, 0-2-0. K, John. 
Taungzun (Thaton), September, 0-45-30. K. John. 
Kinmunsakhan (Thaton), September, 0-3-7, K. John. 
Kyauktan (Hanthawaddy), August, 0-16-0, K, John, 
Syriam (Hanthawaddy), August, 0-2-0, K. John, 
Twante (Hanthawaddy), September, 0-10-0. K. John. 
Kungyangone (Hanthawaddy), September, 0-7-0. K. John, 
Thongwa (Hanthawaddy), August, 3-0-0. Saw San Thwe, 
Dedaye (Pyapon), September, 0-1-0. Maung Ohn Maung, 
Maubin, October, 0-1-0. Maung Ohn Maung. 
Danubyu (Maubin), October, 0-0-1. Maung Ohn Maung. 
Wanetchaung (Insein), September, 0-6-0. K. John, 
Taikkyi (Xnsein), September, 0-4-0, K, John. 
Dam site (Insein), September, 0-5-1. K P John. 
Pegu, August, 0-6-1. K. John, 
Henzada, October, 0-1-0. Maung Ohn Maung, 
Ingabu (Henzada), October, 0-1-1. Maung Ohn Maung. 
Prome, September, 0-4-0. K. John. 
Labu (Prome), September, 0-28-6, K. John. 
Paukkaung (Prome), September, 0-23-2. K. John. 

Pegu Yomas, from 10 miles south of Pyu, September, 0-0-5. G. E. Blackwell. 
Pegu Yomas, from 15 miles south of Pyu (Pegu), September, 0-10-3. G. E. 

Black well. 
Pegu Yomas, from 32 miles south of Toungoo (Toungoo), September, 0-0-1, 

G, E. Blackwell. 
Pegu Yomas, from 24 miles south of Toungoo (Toungoo), September, 0-2-2, 

G. E, Blackwell. 
Pegu Yomas, from 8 miles south of Toungoo (Toungoo), September, 0-1-0, 

G. E. Blackwell. 
Toungoo, October, 0-5-0. K, John, 

Kyaukkyi (Toungoo), October, 0-1-0. Snw Marshall Thwin. 
Sah Der, Karen Hills (Toungoo), September, 0-10. II. I. Marshall. 
Thayetmyo, September, 0-29-7, K. John. 
Tlumbula (Thayetmyo), September, 0-12-2. K. John, 
Magwe, August, 0-10-16, K. John. 





344 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

External characteristics. Size, of smallest, complete and clitel- 
late specimen, 50 by 3 mm. Nephro pores, present from iii, one 
or both of viii usually dorsal, rarely so in ix and then nearer D 
than in viii, apparently always present in x. An equatorial circle 
of tiny white spots is present in each segment from iii posteriorly. 

Spermathecal and male pores, larger than female apertures 
and nephropores, have, in contrast to many other species of 
Drawida, smooth margins and a definite shape. 

A whitened area just in front of and just behind each 
spermathecal and male pore (in x and xi from or just lateral to 
B to or just beyond mBC) is present on every specimen and has 
a definite boundary though not as a rule marked off by a furrow. 
The epidermis of those areas certainly is not thickened and may 
be slightly thinned. Genital markings always are present in the 
whitened areas, Additional markings may be present elsewhere 
(Table 10). 

Internal anatomy. Pigment, in the circular muscle layer, 
lacking underneath white areas near male and spermathecal 
pores but present elsewhere in clitellar segments even though 
unrecognizable externally. Red clitellar coloration, as in other 
drawidas, in the epidermis. 

Esophagus unusually short in the segment behind the last 
gizzard (as in most drawidas) and thence posteriorly wide, with 
thick wall bearing internally shortly papilliform to squarish 
protuberances, closely crowded in longitudinal or vertical rows. 
The vascular plexus appears in cross sections as a thick, dark 
red, uninterrupted layer. Valve, narrow, in hinder portion of 
xxi, anteriorly or posteriorly in xxii. In cross sections the plexus 
is represented by tiny red dots. Tn sections through the proximal 
portion of the intestine the red spots are larger and more closely 
crowded. Gizzards and valves, respectively, of eleven Rangoon 
worms are located as follows: xiv-xv and xxi-xxii (1, intestinal 
origin slightly behind 21/22) ; xiv-xvi, xxi (3, intestinal origin 
at 21/22 apparently) ; xiv-xvi, xxi-xxii (3, intestine beginning 
midsegmentally or posteriorly in xxii) ; xv-xvi (2, intestine be- 
ginning with 21/22 or midsegmentally in xxii) ; xv-xvii (2, in- 
testine apparently beginning with 22/23). Commissures from 
extra-esophag< m K in front of 8/9, Nephridia, present in x. 

Vas deferens, very short, slightly sinuous in 9/10, looped 
around heart of ix and extra-esophageal trunk, almost straight 
in x ? not especially thickened ectally. Prostates, 3-5 mm. long. 



gates: bckmese moniligastimds 345 

Spermathecal atria, 2-3 mm. long, ental half (approximately) 
widened, digitiform, ovoidal or ellipsoidal, abruptly marked off 

from stalk or more gradually narrowed in an eetal portion. 
Stalk wall thick, lumen narrow; more irregular entally. 

Regeneration. Tail regenerate, 6 mm, long and with 40+ 
segments, at 65/66, Substrate length, 22 mm. 

Remarks. Supposed variation in position of spermathecal pores 
relative to 7/8 is slight and may be fictitious, e.g., due to slight 
eversion from the spermathecal duct. 

Intestinal origin was determined in Rangoon worms that had 
heen anesthetized, pinned down straight, and then fixed. The 
esophagus was almost straight. Records of gizzard locations iu 
worms listed above no longer are available. 







Table 1 










Location of 


genital 


markings in 


Z). 


pap\ 


illifer 


pey nana 


Locality 


Segments 

VI) Mil Vlll IX 

I >n i red 
presetal postsetal 


vii viii 
iH(-4lin.n 


Naunggala (*11) 


6 


9 










Taungzuii (45) 


7 


14 






3 




Kyauktan (14) 


5 












Paukkaung (23) 


10 


6 








2 


Tlinton (32) 


7 


6 








1 


Wa notch aung (6) 


6 


2 










Magwe (26) 














Toungoo (4) 












2 


Dam site (4) 














Ingabu (2) 














Labu (32) 












5 


Kinmunsukhan (10) 




1 










Pegu (4) 










1 




Bilin (21) 














Pegu Yomas (22) 




8 






4 


1 


Duyinzeik (22) 


3 


2 










Kungyangonc (7) 














Twante (5) 


1 








2 




Kyaikto (13) 




1 










Auiigsaing (7) 


1 












Tavoy district (12) 






2 









BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Drawida kangoonensis Gates 1925 

Sittang (Thaton), October, 0-1-3. K, John. 

Kyauktan (Hanthawaddy), August, 0-0-2. K. John, 

Rangoon (Hanthawaddy), various dates, 0-5-21. K. John, 

Twante (Hanthawaddy), September, 0-7-9* K* John, 

Pyapon, September, 0-2-1* Maung Ohn Maung. 

Kyaiklat {Pyapon), September, 0-1-4. Maung Ohn Mating. 

Maubin, October, 0-1-3, Maung Ohn Maung. 

Bassein, October, 0-0^3, K. John. 

Pegu, jungle to the east, August, 0-0-2; K, John. 

Myohaung (Mandalay), September, 0-0*14. K. John. 

Bhamo, September, 0-3-14. K + John. 

Myitkyina, September, 0-7-3. K. John. 

External characteristics. Nephropores, present from iii T in 
viii-ix usually slightly dorsal to Z>, apparently lacking in x, 
Clitellar coloration, sometimes lacking in AA of x-xi and more 
rarely of xii 

Spermathecal pores, very small transverse slits exactly on 7/8, 
margins usually smooth but not as straight as in peguana. Male 
pores, very small transverse slits, just in front of or more defi- 
nitely anterior to 10/11, margins almost always slightly tumes- 
cent and often with an appearance of a thickly annular lip. 

A transverse, slightly depressed, translucent area on each side 
of x is in the median half of BC and the equivalent of a setal 
annulus (secondary furrows lacking). The epidermis between 
that area and 10/11 is whitened and rarely with a diagonal or 
longitudinal central area of greyish translucence. Other paired 
genital markings usually are longitudinal but on viii and ix may 
be almost circular or more rarely transverse, in a middle or 
lateral portion of BC of x just lateral to translucent area, 
similarly positioned in ix but in viii more lateral and often just 
median to C ? in viii-ix often in equivalents of setal annuli, in x-xi 
extending across equivalents of presetal and setal annuli and 
often reaching farther posteriorly, occasionally even to posterior 
intersegmental furrow. Median markings, usually between an 
intersegmental furrow and the equator, reach laterally to A, B 
or into BC. All markings except those in setal annulus of x are 
areas of slight epidermal thickening but without clitellar colora- 
tion. 

Internal anatomy. Gizzards, 2-4, in xii-xvii (Table 12). 
Esophagus behind gizzards, usually distended by ingest a, rather 
sigmoid, on the floor al niV one or two longitudinal lamelliform 
ridges, or a horizontal band marked off into 2-4 sections. The 



GATES ; BURMESE MONILIGASTRIDS 



347 



vascular plexus is much thicker, except just behind last gizzard 
and in the valve, than in the intestine. Gizzards and intestinal 
origin, respectively, of Rangoon specimens : xiii-xv, immediately 
behind 22/23 (3); xiv-xvi, midsegmentally in xxiii (1) or im- 
mediate] v behind 23/24 (5) ; xv-xvii, immediately behind 24/25 

(1). 

Commissures from extra-esophageals, in front of 8/9. The 

hearts of viii unite above the esophagus to open into dorsal 
trunk through a single short vertical vessel in median plane. 
Nephridia, of x lacking (10 adults). 

Vas deferens, slender and iridescent en tally, passing ventrally 
median to the heart and then looped around the heart of ix, 
after passing into x in several hairpin loops (not always easy to 
find), then thickened and in a cluster of loops that may be 
nearly as large as or larger than the testis sac. Prostates, 1.0-1,5 
mm. long, capsule slightly widened entally. 

Spermathecal atria, 3-5 mm. long, saccular rather pear-shaped. 
A short, ental portion of the sac occasionally is invaginated into 
the atrial lumen. Spermatozoal iridescence, slight, has been 
noted (several specimens) in atria as well as ampullae. 

Remarks. Nephropores of ix and xi often are difficult to 
identify though probable sites are recognizable even when patent 
apertures were not seen. A more or less pore-like marking oc- 
casionally visible near D of x presumably marks site of a 
nephropore that was functional in juveniles stages. 



Table 11 
Location of genital markings in ZX rcmgoonensis 



Locality 



viii 



ix 



Pal red 



xi 



Segments 

■ ■ ■ 

vi n 



Rangoon 


26 


26 


26 


26 




Kyauktan 


2 


2 


£1 






Pyapon 


2 


2 


3 


3 




Myohaung 


6 


6 


14 


14 


3 


Kyaiklat 


4 


4 


5 


5 


1 


Pegu 






2 


2 


1 


Bassein 


3 


3 


3 


3 




Maub in 


4 


4 


4 


4 




Twante 


11 


13 


16 


16 




Sittang 


2 


3 


3 


3 





Is 



XII 



Median 



2 



Xlll 



1 


25 




2 


1 


2 


1 


13 




5 




2 




1 




4 


3 


13 


3 


1 



Median markings of viii-ix 
Paired marking's are in BG. 



and xii-xiii are postsetal, but in x are presetal. 



348 



BULLETIN : Ml'SEI'M OF COMPARATIVE ZOOLOGY 



Table 12 

Frequency distribution of segmental position and number of 

gizzards in Z). rangoonrnsis 

Segments 



Number of gizzards 



Locality 





■ ■-■ 
-Ml 


■ ■ ■ 

XIII 


\iv 


XV 


x\ i 


xv ii 


Twante (6) 


1 


4 


5 


6 


3 


1 


Maubin (4) 




3 


4 


4 






Pyapon (3) 




3 


3 


3 






Kyaiklat (2) 




2 


2 


9 






Sittaag (1) 




1 


1 


1 






Ky auk tan (2) 




2 


2 


.) 


1 




Bassein (3) 




1 


3 


3 


3 


1 


Rangoon (14 \ 




8 


13 


14 


12 


1 


Totals 


1 


24 


33 


35 


19 


3 



2 



3 



1 

o 



4 
3 
3 
2 

1 
1 
1 

6 
21 



Drawida rara Gates 1925 



4 



f> 



1 

2 

7 
12 



I^amaungthwe River (Tavoy), September, 0-2-0. W. D. Sutton. 

Moulmein (Amherst), August, 0-3-6. October, 0-2-1. K.John. 

Mupun (Amherst), October, 0-26-18. K. John. 

Thaton, September, 0-14-20, K. John. 

Duyinzeik (Thaton }, September, 0-7-0. K. John. 

Naunggala (Thaton), September, 0-0-3. K, John. 

Bilin (Thaton), September, 0-4-0. K. John. 

Aungsaing (Thaton), September, 0-8-5. K. John. 

Sittang (Thaton), October, 0-13-1. K. John. 

Syriam (Ilanthawaddy), September, 0-9-5. K. John. 

Rangoon (Hanthawaddy), various dates, 0-39-36. K. John. 

Kungyaugone (Hanthawaddy), September, 0-2-0. K. John. 

Dedaye (Pyapon), September, 0-1-0. Maung Ohn Maung, 

Bassein, October, 0-15-18. K, John, 

I'amihyu (Maubin), October, 0-2-0. Maung Ohn Mating. 

Hlawga (Insein), September, 0-3-0. K. John. 

Taukkyan (Insein), September, 0-3-1. K. John. 

Ilmawbi (Insein), September, 0-10-5. K. John, 

Wanetchaung (Insein), September, 0-18-3. K, John. 

Taikkyi (Insein), September, 0-23-14. K. John. 

Dam Site (Insein), September, 0-44-2. K. John. 

Pegu, August, 0-16-10. Jungle to the west, 0-14-4. Jungle to the east, 

0-13-10. K. John, 
Tliayetmyo, September, 0-2-0. K. John, 
Allannivo (Thayetmyo), September, 0-2-0. K. John. 
Pyhimana (Yamethiii), October, 0-8-0. K. John. 
Taungdwingyi (Magwe), August, 0-23-18. K. John, 
Magwe, August, 0-4-6. K. Joint. 
Minbu, August, 0-2-2. K. John. 
Alyuhaung (Mandalay), Beptemher, 0-1-4. K. John. 



GATES : BURMESE MQNILIGASTRIDS 349 

Wimtho (Katha), hills to the west, September 0-0-1, Saw San ThwB, 
^Ty itkyiiicL, September, 0-18-14. K. John. 

External characteristics. Length, 35-50 mm, (Taungdwingyi, 
Aungsaing and Magwe), Nephropores, present from iii, close to 
D in viii-ix, apparently always lacking in x. Clitellar colora- 
tion, red, often lacking vent rally in x and a p reset al portion or 
all of xi, the uncolored area often delimited by slightly irregular 
longitudinal furrows at level of lateral margins of male poro- 

phores. 

Spermatheeal and male pores, larger than nephropores and 
certainly larger than female apertures, transversely slit-like. 
Spermatheeal pores, at or just median to C, each usually sur- 
rounded by an annular lip, the pore apparently located on poste- 
rior margin of vii, the lip when specially swollen with an 
appearance of a short transverse papilla. Rarely, a posterior 
portion or even all of the lip is lacking and then the pore appears 
to be on 7/8. The tumescence is believed to be an everted portion 
oi' the spermatheeal duct. 

Genital markings : paired and nearly circular, pre- and post- 
setal in BC of x, transverse and in setal annul i of x and xi just 
lateral to B; impaired, presetal and postsetal in BB (or reaching 
slightly into BC) of viii, x-xi. Markings in setal annul i often are 
slightly smaller than other paired markings, greyish trans- 
lucent rather than white, depressed rather than protuberant and 
without epidermal thickening- Paired markings occasionally pres- 
ent in BC of viii (pre- and postsetal) or vii (postsetal) are 
transverse, indistinctly demarcated areas of epidermal thicken- 
ing, Paired postsetal markings usually (Pyinmana, Taung- 
dwingyi, Minbu, Magwe, Allanmyo, Thatyetmyo) are lacking 
in x. 

Internal anatomy. Gizzards, 2-4, in xii-xviii (Table 13). A 
midventrah typhlosole-like ridge is present in the gut behind the 
gizzards but may not be as obvious as in rangoonensis. Location 
of gizzards and intestinal origin, respectively, in several Rangoon 
specimens: xiv-xvi, midsegmentally in xxii (3) or just behind 
22/23 (2) ; xv-xvii, just behind 22/23 (4) or midsegmentally in 
xxiii (12) or just in front of 23/24 (1) or just behind 23/24 (2) ; 
xv-xviii, midsegmentally in xxiii (2) ; xvi-xvii, just behind 22/23 
(1) or midsegmentally in xxiii (1) or just behind 23/24 (2) ; 
xvi-xviii, just in front of 23/24 (1) or just behind 23/24 (6). 
The intestine begins, in a Taungdwingyi worm with gizzards in 
xiv-xvi, just behind 21/22. Commissures from extra-esophageals, 
in front of 8/9. 



350 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Vas deferens, short, slender, with several loops in fl/10 and 
on anterior face of 9/10, twisted a round hearts of ix, with several 
further loops on posterior face of 9/10 in a small compact cluster, 
an eetal portion only slightly thickened. 

Abnormality. (No. 1.) Right spermathecal duet, about half 
way down to parietes, passes into vii where a short portion is 
widened into a thin-walled vesicle. (No. 2,) Right male poro- 
phore entirely in x and extending from the presetal secondary 
furrow nearly to 10/11. The male pore is about midway between 
eq/x and 10/11. (No, 3.) Testis sac, vas deferens, prostate, 
male pore and porophore as well as genital markings of left 
side, lacking. 

Regeneration. Tail regenerates, of 28+ segments at 72/73 and 
of 26+ segments at 81/82. Lengths of substrates and regenerates, 
respectively, 28 and 4, 31 and 5 mm. 

Remarks, Male porophores of four Taikkyi worms are repre- 
sented only by circular areas of epidermal whitening, distinctly 
demarcated but not protuberant. The pores are transversely 
placed slits at 10/11 which appear to be nearly as deep across 
the porophore as elsewhere. Prostates of these specimens are 
unusually high in the coelomic cavity though capsules still are 
almost confined to the parietes. 

Male porophores in vara seem incapable of marked elevation. 
However, by bringing the prostatic capsule deeper into the 
parietes, a small part of the porophore containing the male 
aperture can be raised into a teat-like protuberance presumably 
for insertion into the spermathecal pore. The lumen in the 
thickened portion of the spermathecal duet may be large enough 
to receive the protuberance. Entrance perhaps is facilitated by 
slight eversion of the duct. 

The unusually small size of the prostatic capsule presumably is 
secondary rather than primitive but in either case distinguishes 
vara from eonsiricta Gates 1929. Connectives from the extra- 
esophageal trunks are on opposite sides of septum 8/9 in the 
two species. Discoidal male porophores extending into x and xi 
have been developed in both species but seem capable of greater 
elevation (as temporary intromittent organs) in eonsiricta. 
Genital markings of both species are similar to those of spissata 
Gates 1930 where they also are not associated with definite glands. 
The male pores of spissata certainly do appear to be well inside 
xi but all specimens had protruded porophores (discoidal on re- 
laxation and restricted to xi-). Assuming that adivertieulate 



GATES: BURMESE MONILTGASTRTDS 



351 



spermathecae are evidence, in Burma, of phylogenetic relation- 
ships, spissatA can be derived from a common ancestor of vara 
and constrict a by the following changes: migration of male 
pores back into xi, increase in number of gizzards (now ap- 
parently 4-5), elongation of esophagus so as to place the gizzards 
farther posteriorly, elongation of prostates as well as male 
deferent and spermatheeal ducts. Additionally, the spermathecal 
duets have been thickly museularized, especially so in the ectal 
35-50 mm., and the vasa deferentia (in x only) have been 
thickened. 



Table 13 
Frequency distribution of segmental position and number of 

gizzards in D, rant 

Segments Number of gizaards 

Locality 





* * 

Ml 


XIII 


xiv 


XV 


xvi 


\ v 1 i 


i ■ ■ 

xviii 


if ague (15) 


1 


12 


15 


15 


9 






W imt ho (2) 




1 


1 


2 


»» 






Ta-imgthvingyi (6) 




1 


5 


6 


4 






Tim ton (11) 




1 


11 


11 


11 


8 




Pegu (22) 




6 


17 


22 


g.g 


14 


2 


Mupun (15) 




2 


12 


15 


15 


10 


2 


Myoliaung (5) 






4 


5 


5 


1 




Taikkyi (5) 






4 


5 


5 


1 


1 


Rangoon (9) 






1 


9 


9 


6 


2 


Bassein (2) 








2 


£ 


2 




Syria ni (2) 








2 




2 


2 


Totals 


1 


23 


70 


94 


86 


44 


11 



2 


3 


4 




8 


7 


1 




1 


2 


4 






2 


9 




5 


17 




4 


11 




5 






4 


1 


2 


5 


2 
2 
2 


5 


37 





Drawida tenellula nom. nov. 

1933. Dru-irida sp„ (rates, Ret-. Indian Mus., 35:470. 

Remarks. Ovisacs were small in both specimens (aclitellate) 
but other genital organs probably were fully developed. 

The "penes," if definite structures, presumably are retractile 
into chambers possibly restricted to the parietes. Otherwise, the 
penes could be produced perhaps by e version of the prostatic 
duct. In either case complete retraction is likely to leave a fairly 
large secondary male pore that would provide an additional 
distinction from the various species mentioned below, Pro- 
trusion of a discoidal porophore in a tubular fashion now seems 
unlikely and is contra-indicated by texture of penial epidermis. 

D. tenellula is distinguishable from bullata by the penis-like 
male porophores and absence of definite genital markings. From 




1152 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

other species with shortly digitiform spermathecal atria D, 
tenrlhtla is further distinguished as follows. From D. vulgaris 
by the longer vas deferens. From ZJ. gracilis by the much 
smaller size, smaller ventral setae of iiivii, erect prostates, 
thickening of the vas in xi, peripheral closure of ovarian chamber. 
From 2). ftexa by the longer vasa deferentia and absence of 
special parietal glands associated with genital markings. 

Drawida Victorian a n. sp. 

Mount Victoria (Pakokku Chin Hills), on east side near path from Knupet- 
let to summit, at 3000 feet, in tenk forest and grass hind, July, 0-1-0. 
G. Heinrieh. 

External characteristics. Length, 64 mm. Diameter, 3 mm. 
Setae, unrecognizable or almost so in an anterior portion of the 
body, posteriorly AA < BC\ AB slightly < CD, Nephropores, 
present from iii, at or close to D. 

Spermathecal apertures, probably at 7/8 and at or just median 
to C, Male pores, in x and very slightly lateral to B y very small 
transverse slits at tips of anteriorly directed, rather conical 
protuberances from posterior margin of x. Porophores are in 
slight depressions where the epidermis is whitened, smooth and 
glistening. 

Genital markings, small, median and postsetal in vii-xi, paired, 
in BC and about in line with male porophores, in presetal and 
also in setal annulus of x, in postsetal annulns of xi, A smaller 
presetal marking is present in lateral portion of A A in x. Each 
marking, except in vii, nearly circular, with a distinctly de- 
marcated rim and a longitudinal or transverse area of greyish 
transl uccnce centrally. 

Internal anatomy. Septa, 5/6-8/9 muscular. Gizzards, four, 
in xvi-xix. Commissures from extra-esophageals, in front of 8/9, 

Testis sues, dislocated posteriorly behind the ovarian chamber. 
Vas deferens, slender and iridescent entally in 9/10 and in ix 
near hearts, a preseptal portion in ix 5-10 mm. long and in a 
small cluster of u-shaped loops, the postseptal portion thickened 
and in a cluster of loops that is as large as or larger than the 
testis sac, passing directly into prostate just below its ental end. 
Prostates, rather slenderly club-shaped, 2y 2 mm. long, nearly 
straight or bent into a u-shape, capsule slenderly club-shaped 
and nearly circular in cross section. 



GATES: BURMESE M0NILIGASTR1DS 353 

Spermatheeal duet, slender; slightly zig-zag looped as it passes 
ectally on posterior face of 7/8. Atrium, saccular, large, erect in 
vii and in contact with dorsal parietes, the octal half stalk-like. 
Annular ridges are present on inner wall of stalk and sac. 
Ovarian chamber, probably closed off from parietes. Ovisacs, in 
xii-xiv or xv. 

Longitudinal musculature, uninterrupted over sites of genital 
markings. 

Remarks. Pigment was not recognized and may be lacking but 
the body wall had been deeply stained by some post-mortem 
accident. The anterior end was distorted probably as a result 
of local desiccation. 

Yolk was present only in a posterior portion of partially col- 
lapsed ovisacs, a condition that is suggestive of an early post- 
sexual stage. The glandular investment of the prostates, thin 
and firmly adherent to the capsule, is more like conditions of a 

presexual stage. 

Genital markings of the kind present on the type, in other 
species usually are associated with small solid glands having 
transparent walls. If such glands are present, vietoriana is dis- 
tinguished from papillifer and nag ana by the protuberance of 
the male porophores and from nepalensis by location of male 
pores in x as well as by characters of the porophores. If glands 
are absent, vietoriana is distinguished from rangoonensis by the 
protrusibility of the male porophores and from molest a by absence 
of penes. To permit entry in a key, glands are assumed to be 

present. 

Reproduction: Iridescence in an ental portion of the male 
gouoduets shows that sperm had been matured, A similar irides- 
cence in the coagulum of an ectal portion of the atrial stalk 
shows that sperm had been received, presumably from another 
worm. Reproduction, accordingly, is assumed to be sexual and 
biparental. (No iridescence was recognizable in the coagulum 
within the spermatheeal ampulla.) 

Regeneration. A tail regenerate is 3 mm. long, 

Drawida vulgaris Gates 1930 

Frame, September, 0-2-12. K. John. 

Paukkaung (Prome), September, 0-18-20, K. John. 

Mt. Popa (Myingyan), September, 0-0-2, K. John. 

External characteristics. Length, to 50 mm. Diameter, 2%-S 
mm. Nephropores, present from iii, in or close to CD, apparently 



354 BULLETIN ; MUSEUM OF COMPARATIVE ZOOLOGY 

lacking in x. Setae, A A usually < BC but on some Paukkaung 
worms may = BC f DD ca. = ^C. 

Male pores, small transverse slits on posterior margin of x, at 
or slightly median to mBC, with or without an obvious though 
small, annular lip. A posterior portion of x bearing the male 
pore may be protuberant as on the types or not. In the latter 

case the male pore is concealed from view by the anterior margin 

£■ 
XL 

Genital markings, transverse, shortly elliptical, presetal, opaque 
areas, each with a greyish translucent center, in Paukkaung 
worms indistinctly delimited and recognizable only in best- 
optical conditions. Paired markings usually are in a lateral 
portion of BC, with centers slightly lateral to levels of male 
pores, A pair of whitened patches usually slightly nearer B on 
each of the present worms extends from 10/11 to postsetal 
secondary furrow of x. In one or the other of those patches there 
occasionally is recognizable a typical marking with central trans- 
lucence. Markings apparently are lacking on the Popa worms. 
The markings of Paukkaung and Prome worms, respectively, 
are as follows: in ix (88 and 12 specimens), in x (38 and 12) ; 
median, in AA, of viii (1 and 5) ; of ix (17 and 9) ; of x (0 
and 4) ; of xi (19 and 7) ; of xii (14 and 6) ; of xiii (1 and 0). 

Internal anatomy. (Jizzards, two to four in xii-xvi, xii-xiv (3 
Prome), xii-xv (1 Paukkaung), xiii-xiv (1 Paukkaung, 2 Prome ), 
xiii-xv (4 Paukkaung, 2 Prome), xiv-xvi (1 Popa). Commissures 
from extra-esophageals, in front of 8/9. Posterior commissures 
from extra-esophageals pass into hearts of ix just lateral to 
median plane. Hearts of ix unite mesially just above gut and are 
then connected with the dorsal trunk by a short, vertical vessel 
in the median plane* A vessel from the posterior commissures 
of the extra-esophageals passes posteriorly on dorsolateral face 
of the gut to xii or xL Nephridia of x, lacking. 

Testis sacs, usually laterally flattened, equally in ix and x, 
rather reniform to shortly u-shaped, concave ventrally, more or 
less deeply constricted by 9/10. Vas deferens, slender and 
iridescent entally, passing into ix median to the heart, then 
anterior to the heart and back to 9/10 where it is twisted into 
several short loops, the portion in x thickened and opaque, also 
twisted into several loops, then passing directly into the prostate. 
The vas of one Prome worm is about 8 mm, long, length of the 
slender portion just over 3 mm. Prostatic capsule, about one 
mm, long, slenderly rod-like but with some slight ental widening, 



GATES: BURMESE HON ILIGASTRIDS 355 

soft, opaque, of about tin* same thickness as the glandular in- 
vestment. 

Spermatheeal atria, always in vii, about one mm. long. 

Remarks, Differences from hull a fa are few but apparently 
constant in a range that extends into Thailand. Vasa deferentia 
of all specimens referred to vulgaris, like the spermatheeal atria, 
have been short as in the smallest specimens of bullata. Male 
pores of bull at a always were in or appeared to be in xL Tn 
nulgaris, however, the male pores always were in or appeared to 
be in x, 

Table 14 
Location of genital markings in ZX vulgaris 

Segments 
taacaiity ix x viii i\ x xi xii xiii 



Paired Median 

Paukkatmg 38 38 1 17 — 19 14 1 

Prome 12 12 5 9 4 7 6 — 

Paired mnrkingg are located laterally in BC\ Median markings are confined 
to A A, 

Table 15 

Frequency distribution of segmental position and number of 

gizzards in D. vulgaris 

Segments Number of gizzards 

\ii xiii xiv xv xvi 2 3 4 



Locality 



1 


4 


2 


5 




1 


3 


10 



Prome (6) 3 7 7 2 

Paukkamig (7) 1 6 6 5 
Mt. Popa (1) 111 

Totals 4 13 14 8 1 

Genus MOXTLIG ASTER Perrier 1872 

MO N1L1G ASTER CERNOSVITOVI 110111. 11 OV. 

1940, Moniligasfer beddardi Gates, Bee. Indian Mus., 42:496. (Type lo- 
cality, KodaikanaL Types in the Indian Mus. ) 

M. beddardi Gates 1940 is preoccupied by M, beddardii Rosa 
1890 now Drawida beddardi, and would have been renamed long 
ago had there been any occasion to refer to the genus. 

MQNILIGASTRID PHYLOGENY 

The numerous differences from most earthworms suggest long 
separation of moiiiligastrid stocks from other lines of megadrile 



356 BULLETIN : MUSEtM OF COMPARATIVE ZOOLOGY 

evolution. Unless peculiar characteristics, in identical combina- 
tions, evolved independently in the ancestry of each genus, an 
early moniligastrid should have differed from contemporaneous 
oligochaetes as follows: prostomium, now independent of the 
peristomium and attached to the roof of the buccal cavity behind 
level of intersegmental furrow 1/2; digestive system, with en- 
teroscsjmental organs, with esophagus elongated posteriorly so 
that the gizzard is behind the ovarian segment, perhaps with 
intestinal origin at first in xv where it still is today in many 
megadriles; vascular system, with paired extra-esophageal trunks 
that are lateral to the hearts, with a parietal subneural trunk, 
one or two pairs of hearts at posterior end of the series united 
dorsomesially in each segment instead of opening into the dorsal 
vessel directly; testes, proliferating anteriorly into interior of 
their septa and no longer inducing development of seminal 
vesicles ; ovarian chamber, closed off mesially from neural and 
esophageal portions of the coelom; spermathecae, deeply in- 
vaginated into the coelom so that ampullae are dorsal ; capsular 
genital glands, evolved from the X glands. While so much 
typically moniligastrid anatomy w r as being evolved, gonads of 
the ancestral battery may well have been eliminated in all but 
four segments, x-xiv. 

Ancestral characters still retained were as follows : genital 
apertures, of all organs from spermathecae to glands, minute, 
superficial, in the ventrum; spermathecal, male and female pores 
in region of AB; gland pores variously located in BB: sigmoid 
setae, in four pairs of longitudinal ranks; clitellum, unilayered; 
gonoducts short, and opening directly to the exterior in preset al 
portions of the segments; hearts, lateral (a supra-esophageal 
trunk still lacking); ovisacs, dorsal and posteriorly elongated; 
gizzard, still esophageal and single ; nephridial ducts, passing 
straight through the parietes. Pigment may have been lacking. 

An important early change in moniligastrid anatomy resulted 
from acquisition by genital glands of ability to attract toward 
and into themselves during early growth the elongating male 
gonoducts. At first gonoducts may have joined, regardless of 
location^ the nearest glands, or if several were equidistant those 
that were growing faster or had invaginated earlier. Eventually 
stabilization was achieved and the vasa deferentia joined only 
those glands developing at the intersegmental level next behind 
that of the gonad septum. Henceforth, male gonoducts were to be 
unable to open to the exterior in absence of prostates. 



GATES: BURMESE MONILJGASTRIDS 357 

Reduction in the gonad battery was continued. In a line 
leading to Desmogaster, gonads of x were aborted and those 
of xii became testes. In lines leading to Hastirogaster and 
Eupolygaster, gonads of x and xii, of x and xiii, respectively, 
were eliminated. In the main line leading to Drawida-Monili- 
gaster, gonads of xli-xiii disappeared (unless a segment was 
exealated anteriorly), those of xi being retained as ovaries. 
Other reductions and modifications in the battery would seem to 
have been possible. If so, additional genera may be found in 
unexplored portions of the moniligastrid domain east of Burma* 

Development of additional hearts at posterior end of an 
ancestral series appears to have been limited by location of the 
ovarian chamber as none have appeared behind a level two seg- 
ments in front of that chamber. Thus, the last hearts of drawidas 
arc in ix but of desmogasters are in xi, One pair of spermathecae 
was eliminated in the ancestry of Drawida (and Hastirogaster?) 
but elsewhere the reduction has been intrageneric. 

Subsequent to appearance of extant genera, known macro- 
scopic changes have been confined almost entirely to short por- 
tions of nephridial ducts, a section of the gut behind the ovarian 
segment and, of course, to the genital system. Parietal portions 
of nephridial ducts now may grow laterally as well as mesially 
within the body wall to acquire at several levels external open- 
ings, often irregularly placed and with more or less asymmetry. 
The esophagus has been further elongated. Esophageal gizzards 
(cl Tables 16-17) have been multiplied and moved posteriorly. 
Body size has been increased sufficiently, in various countries, 
to warrant the characterization of "giant earthworms/ 7 These 
somatic modifications now appear to have been made independ- 
ently in various genera. 

Among independently made changes in the genital system are: 
lateral dislocations of male and spermathecal pores; closure of 
the ovarian chamber from the parietes; elongation of sper- 
mathecal and male ducts; development of intromittent organs; 
various modifications in shape of the prostatic capsule as well 
as reduction (and final elimination?) of the glandular invest- 
ment. 

Female pores have remained in their primitive position 
throughout the family, and spermathecae still are adiverticulate 
except in various lineages of Draiuida. Although more informa- 
tion is available for that genus than for all others it is not yet 
sufficient to allow more than a tentative indication of relation- 
ships and changes in a few small groups. 



358 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

A rather primitive willsi species group has retained prostate- 
like (GM) glands but has acquired pigment. Spennathecae still 
are adiverticulate in the Assamese limctta Gates, 1934, but pores 
are at mJSO, gizzards are three in xiii-xv and the prostatic capsule 
has become so small that it is almost confined to the parietes. 
Genital apertures still are at or close to B in species of penin- 
sular India which now have spermathecal atria. D. scandens 
Rao, 1921, with prostates and GM glands of the same size, lias 
two or three gizzards in xii-xv. I). dolosa Gates, 1945, with two 
to four gizzards in xiii-xvi, has shortened prostates that may be 
only a quarter as long as the GM glands. Z). period iosa Gates, 
1934, with two or three gizzards in xiii-xvi has prostatic capsules 
so small as to be almost confined to the bodv Avail. D. willsi 
Miehaelsen, 1907, with two to four gizzards in xii-xvi, has GM 
glands that are shorter and slenderer than the prostates. Occa- 
sional individuals have no glands. 

All Burmese species have spermathecal pores at or close to 
C and a. region around site of each primary male pore usually 
is modified to some sort of a porophore for more efficient transfer 
of sperm during copulation, 

A beddardi group, probably with a peripherally closed ovar- 
ian chamber, lacks pigment and spermathecal diverticula 
though a terminal portion of the spermathecal duct is thickened. 
D. delicata, with four gizzards in xii-xv, has markedly lengthened 
male gonoducts but spermathecal ducts still are short, D, hempi 
Stephenson, 1914, with gizzards in xv-xix, has spheroidal pro- 
static capsules. D. vara, with three to five gizzards, in xii-xviii, 
has prostatic capsules that are vestigial and confined to the 
parietes. Z), constricta, probably close to ram but with only two 
or three gizzards in xiv-xviii, has less shortened prostatic cap- 
sules that still reach into coelomic cavities. 7). spissata Gates, 
1930, with four or five gizzards in xvii-xxi, has much lengthened 
prostates, vasa deferentia and spermathecal ducts, the latter 
markedly muscularized ectally. D. beddardi, with three or four 
gizzards in xii-xix, still has digit if orm prostatic capsules but 
with the glandular investment occasionally restricted to small 
ental portions, Male pores have been invaginated and now are 
on tubular penes pendent from roofs of eversible, muscular- 
walled chambers more or less protuberant into the coeloin. 

Possibly derived from a distant ancestor of the beddardi 
group is caerulea, with pigment, male pores deeply invaginated 



GATES: BURMESE HONILIGASTRIDS 359 

into muscular copulatory chambers that have no glandular in- 
vestment and contain no penes, fipermatheeae, in the southern 
portion of the range, still are adiverticulate though the terminal 
thickening of the duct may be slightly larger than in the 
heddardi group, A muscularized atrial outgrowth of the thick- 
ened portion of the duct has appeared in more northern forms 
and still farther north has reached septum 6/7 and the dorsal 
parietes. Elongations already under way in the north have 
become even greater in the Assamese decourcyi with atria, sper- 
mathecal and male ducts, respectively, reaching lengths of 8, 
20 and 650 mm. Gizzards, still onlv 1 or 2 in xiii-xvi of a central 
Burma population, in decourcyi now are 7-9 in xiv-xxvii. 

Other Burmese species, all with diverticulate sperm athecae, 
appear to belong in two groups of more or less closely related 
forms distinguished by presence or absence of definite glands 
(not prostate-like) in association with the genital markings. One 
section of the ^landless group has digit if orm spermathecal atria. 
Three species still have digitiform prostatic capsules: Z). vulgaris, 
with 2-4 gizzards in xii-xvi, short atria and vasa deferentia; D. 
tenellula, with 3-4 gizzards in xiii-xviii, and lengthened vasa; D. 
bitttata, with 1-5 gizzards in xiii-xviii, atria and vasa that vary 
from short to considerably elongated. /). (jracilis* with 1-4 giz- 
zards in xiv-xxi, short atria, lengthened vasa but prostatic cap- 
sules that reach into coelom only slightly. 7). Imeriom, with 3-5 
gizzards in xiii-xx, short but muscularized spermathecal ducts 
and atria, has lengthened vasa, anteroposterior^ elongated and 
bilobed prostatic capsules sessile on the parietes. 

An ental portion of the somewhat elongated spermathecal 
atria, in the second section of the glandless group, has become 
a thin-walled sac. D. rangoovensis, with 2-4 gizzards in xii-xvii, 
has elongated vasa but nearly digit if orm prostatic capsules, D. 
molesto, with 3-5 gizzards in xiii-xxi, elongated vasa and pro- 
static capsules, has invaginate male pores on tubular penes. 

<iM glands of the remaining group are solid and bear on their 
outer faces part or all of a more or less distinctly delimited gen- 
ital marking. Most species have digitiform atria. Two of the 
species in that section have clear glands with transparent or 
translucent walls. D. asmmensis, with 4-5 gizzards in xii-xvii, 
has short spermathecal ducts, atria, vasa deferentia and prostatic 
capsules. D. ftexa, in peninsular Burma, also has short atria 
and vasa but the gizzards (4-5) are in xvi-xxiv and glands are 
much more numerous. D. nana, with 2-3 gizzards in xiii-xv, has 




36f) 



BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 



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362 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

glands with opaque bill soft walls. Vasa, atria, possibly sper- 
matheeal duets and prostates, are lengthened. The rest of the 
species in the section have glands with thickly muscular walls. 
/), tumida, with 3-5 gizzards in xvi-xxii, has short atria but vasa 
are somewhat elongated. D. longatria, with 1-6 gizzards in xiii- 
xxiii, has spennathecal atria 2-180 mm. long, Male gonoduets 
are of similar variable lengths. 

An ental portion of the somewhat elongated spennathecal atria 
has become a thin-walled sac in other Burmese species which have 
almost digitiform prostatic capsules and (one exception?) clear 
glands. D. papillifer, with 2-5 gizzards in xii-xx still has short 
vasa, no protuberant male porophores and an ovarian chamber 
that is not closed off from the parietes. Pigment, possibly 
similar to that of eaemlea, is present. Z>. nepalensis and vic- 
toria no, with gizzards in xii-xxiii and xvi-xx, respectively, now 
have elongated vasa, protrusible male porophores and an ovarian 
chamber closed off from the parietes. 

OLIGOCIIAETE PIIYLOGENY 

The supposedly Jurassic or Cretaceous ancestor of the nieg- 
adriles usually was believed to be octogoiiadal, with testes in 
x-xi and ovaries in xii-xiii. Divergent conditions in modern 
earthworms, except moniligastrids, resulted from elimination of 
the anterior ovaries and, less frequently, of the first or second 
pair of testes. The phylogenetic puzzle posed by the Monili- 
gastridae required answers to several questions among which 
were the following: 1) How had testes of x-xi gotten into xi-xii 
during evolution of Desmogastrr from the supposedly ancestral 
liaplotaxicl? 2) How did ovaries of xiii get into xi as Drawida 3 
was evolving from Desmogoster? 3) At the same time, how did 
one pair of dcsmogaster testes, either from xi or from xii, get 
into x? 4) How had male gonoduct funnels gotten from one 
septum into sacs of the preceding septum? 

Evolution of gonadal batteries within the Moniligastridae re- 
quires "some possibility of the intercalation or excalation of seg- 
ments' * according to Beddard (1891) who offered no suggestions 
as to how the deletions or insertions could have been brought 
about. The only attempt at a solution of the moniligastrid puzzle 
is Stephenson's ingenious "contraction" theory which Avel 



8 Drawida is understood, in most of tliis discussion, to include Mnniligastrr. 
The genera are sr> much alike that Michuelsen seems to have repented separating 
them. Mtniiliffnstcr mat be only an unusually distinct species group. 



GATES: BURMESE MONILIGASTRIDS 363 

(1959) deems plausible. Michaelsen, however, appears to have 
been unwilling to approve — except for mention of several diffi- 
culties in a footnote (1922), he continued to ignore the problem 
which certainly was involved in his own derivation of the Monili- 
gastridac from the Haplotaxidae. 

If moniligastrid testis sacs, as Beddard suggested, are vestigial 
eoeloms of segments from which all parts except gonads, gono- 
ducts and their funnels have disappeared, it should be possible 
to determine the ancestral condition by li expanding" the sacs 
to their original metamerie state. The genera thus theoretically 
treated by Stephenson are Syngenodrilm and Dcsmogaster. Re- 
sults were identical, testes in xii as well as x and ovaries in xiv 
(for figures cf. Stephenson, 1922 or 1930). Then, to restore the 
primitive continuity of the series, testes were inserted in xi and 
ovaries in xiii thus providing a deeagonadal battery. Conditions 
in most megadriles resulted from elimination of gonads in xiv, 
xii and x or xi but in moniligastrids resulted from partial or 
complete abortion of gonad segments. Contraction of the tenth 
and twelfth segments to testis sacs along with elimination of 
testes in xi and of ovaries in xiii produced Desmogaster. Fusion 
of desmogaster testis sacs and elimination of an intervening seg- 
ment to bring testes of x into association with male funnels 
formerly in xii, gave Draw i da. 

Various sorts of evidence^ including some from SyngenodriluB^ 
were cited in support of the contraction theory, 1) The epithelium 
lining the moniligastrid testis sacs requires the cavity of the sac to 
be coelomic. 2) Trabeculae, within the sac ? were thought to be 
remnants of former septa, 3) The ovarian chamber of various 
moniligastrids has only to be separated into discrete halves along 
with loss of nephridia to reproduce identically the testis sac 
condition, 4) Similar contractions are said to be under way in 
various megadrile families. 

Although Miehaelsen's criticisms now appear unimportant, 
contraction still is unacceptable and for the following reasons. 
1 ) Expansion of Syngenodrilits testis sacs involved misinterpre- 
t at ion of figures and text of an inadequate and, in part, 
erroneous characterization of the type species, Independent 
examinations of the type (Gates 1945, and Pickford, 1945) 
found testis sacs to be membranous enclosures of small portions 
of coelomic cavities. These sacs are in no way comparable to 
the so-called testis sacs of moniligastrids and there had been 
no reduction in size of testis segments, 2) Desmogaster gonads 



364 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

were assumed to be one segment in front of the locations men- 
tioned in the literature 1 . Subsequent investigations have con- 
firmed the original records. 3) Evolution of the moniligastrid 
ovarian chamber has involved no reduction in size of the 
ovarian metamere. Essentially, all that has happened is a 
partitioning off from the rest of the eoelomie cavity of a por- 
tion containing the ovaries, nephridia and oviduct funnels. 
Halving the sac and eliminating nephridia would leave size of 
ovarian segment unchanged and would not provide sacs homolo- 
gous with those containing the testes. 4) Similarly, supposed 
contractions in Acanthodrilus, Hoplochaeiella and Eutyphoeus^ 
as well as various changes in other genera, all involve sub- 
division of eoelomie cavities but always without any real de- 
crease in volume of eoelomie space or in size of metaraeres. 5) 
Moniligastrid testis sacs (ef. p. 301) have no real cavity and no 
internal epithelium aside from the male funnel. Trabeeulae 
are only muscle strands without peritoneal covering, 

The moniligastrid problem was made unnecessarily difficult 
by th<> phylogenetieally-based classical system in which Drawida 
is directly descended from Desmogmter. If Eohippus could not 
remain Eohippus during all of the time it was evolving into 
E quits, Desmogaster is unlikely to have remained Desmogaster 
while evolving into Drawida. Hence Drawida ovaries of xi and 
Desmogaster testes of xii, as w 7 ell as other aspects of the puzzle, 
can be considered separately and independently rather than 
simultaneously. 

The Drawida condition can be derived theoretically from the 
ancestral octogonadal battery by elimination of the ovaries in 
xiii and the testes of x along with excalation of one segment 
in front of the gonad region. The gonads thus eliminated did 
disappear in evolution of the Enchytraeidae — if that family has 
a common ancestry with megadriles. The required excalation has 
been made (ef. below) in another family. 

The Desmogaster condition cannot be derived by any of the 
evolutionary changes recognized in the classical system. Here 
also the problem was made unnecessarily difficult by an assump- 
tion of gonad immutability. Although never so stated, construc- 
tion of the basic phylogeny proceeded as if the sexual nature of 
each gonad in the ancestral battery already had been irreversibly 
determined. Studies of partheiiogenetic morphs by the author 
have shown that any testis can become hermaphroditic or even 
an ovary of different and distinctive shape. Determination of 



GATES ; BURMESE MONILIGASTRffiS 365 

gonads for maleness accordingly is reversible. Although that 
discovery is of no help in solving the Desmogaster problem, it 
does provide a second method of deriving the Drawida condition 
from the octogonadal battery, viz., by elimination of gonads in 
xii and xiii along with conversion of testes in xi to ovaries. 
Conversion of ovaries to testes has not been found in the author's 
morphs and is not to be expected in series hitherto studied, as 
oligochaetes seem to have an innate prohibition against evolution 
of dioecism, 4 The literature does, however, contain various rec- 
ords of gonads in xii of normally mctagynous species — those in 
which the gonads of xii usually abort ontogenetically. Unaborted 
gonads of xii in such individuals were ovaries, hermaphroditic, 
or even testes. The Desmogaster condition then can be derived 
merely by eliminating from the ancestral octogonadal battery the 
testes in x and bv conversion of ovaries in xii to testes. 

Gonadal batteries of two megascolecid genera, Tonoscolex and 
Nello$eole& Gates 1933 and 1939, with testes in ix-x and ovaries 
in xii, obviously can be derived only by excalation of one meta- 
mere in front of the gonad region. Fortunately, somatic as well 
as reproductive organs from the gizzard and posteriorly are 
one segment anterior to normal location. The exealated segment 
must then have been one of the first four or five. Just how the 
excision could have taken place is shown by changes now under 
way or recently made in various megadriles. The intersegmental 
furrow demarcating the first from the second segment, even 
when preservation is optimal, often is indistinct. That same 
furrow, along with the prostomium, has disappeared in Pon- 
toscolex corethrurus (Miiller, 1856) and the first two segments 
have become so small that together they are markedly shorter 
than the third. Actually, only presence of setae just in front of 
the first intersegmental furrow (originally 2/3) enables recog- 
nition of the fusion, just as in the apparent peristomium of the 
leech-like oligoehaete (cf, above p, 298), Acanthobdella peledina 
Grube, 1851, and in Trichodrilus caniabrigiensis (Beddard, 
1908). Elimination of the setal follicles originally belonging in 
ii, that is now under way in various species, when completed 
will produce the elision required by Tonoscolex and Nelloscolex. 

Also noteworthy in the same connection is the derogation of 
metameric organization anteriorly that has become common, pre- 
sumably subsequent to retraction of the brain into the region of 

4 All supposed eases of dioecism now have been found to be instances of genital 
polymorphism in which the morphs have evolved after establishment of partheno- 
genesis. 



306 BULLETIN ; MUSEUM OF COMPARATIVE ZOOLOGY 

segment iii and of the united cord ganglia of i-iii to a slightly 
more posterior level. Among those long known "cephalizations" 
are various abortions, presumably often ontogenetic : of setal 
follicles in ii as well, sometimes, as several successive segments; 
of portions or all of one or more of septa 1/2-4/5; of nephridia 
in ii or occasionally even in additional segments as far back as 
x-xiv; of parts or all of vascular commissures between dorsal 
and ventral trunks in ii-iv or v, of the dorsal trunk itself and 
back to hearts of vi, vii, viii or even ix. 

Escalation of one segment, along with elimination from the 
octogonadal battery of one pair of ovaries arid/or one pair of 
testes provides gonadal conditions in most microdriles. Two 
exceptional lumbriculid genera, Sfyloscolex and Premnodrilits, 
require two segments to have been elided. Even for that deriva- 
tion evidence is provided by location of somatic as well as genital 
organs in an unnamed glossoseolecid (Gates, MS) at least two, 
possibly three, segments in front of the usual positions. 

Gonadal conditions in the few remaining microdriles, Aeoloso- 
matidae and Naididae, can be derived theoretically from the 
ancestral battery by elisions of 4, 6 or 7 segments. For so much 
excalation no evidence has been found* A different way of 
bringing naids into a common line of oligoehaete descent requires 
further consideration of the genital conservatism that is basic 
in the classical system. 

The only changes allowed in the gonadal series during the 
entire period of megadrile evolution are elimination of a pair 
of testes and/or of one pair of ovaries. The author's studies of 
parthenogenesis have shown that all reproductive organs, except 
the elitelluiii, ovaries and female gonoclucts, can disappear in 
such a short time as to allow no evolutionary changes in spe- 
cifically distinctive, somatic structure, Although genital evolu- 
tion presumably is slower in sexually reproducing forms, 
opportunity for many more than the one or two gonad elimina- 
tions of the classical system does seem to have been provided 
during the Tertiary and Quaternary. Former existence of a 
longer ancestral series, with gonads at least in segments v-xviii, 
now is suggested because of the following facts: 1) Location of 
ovaries in two species of Haplotaxis as Tar bark as xv-xvi, 
gonads formerly in xii-xiv presumably eliminated* 2) Occa- 
sional presence in lumbricid individuals of ovaries in some or 
all of xiv-xviii. 3) Occasional presence of ovaries in xiv of 
individuals belonging to four other families. 4) Presence of 
gonad anlage in xii of embryos, various species (3 families) with 



GATES: BURMESE MONILIGASTRIDS 367 

no gonads in that segment of adults. 5) Presence of testes in ix 
of occasional ociierodrilid and niicroehaetid individuals, 6) 
Presents ol' testes in some or all of segments v-ix, all individuals 
of one common, parthenogenetie morph of Pheretima anomala 
Michaelsen, 1907 (cf. Gates, 1956), of especial interest in con- 
nection with the Naididae where testes are in one of segments 
iv-Yii. 7) Gonads in as many as nine segments of head regen- 
erates, several species of Perionyx. 8) Gonads in as many as 
twelve segments of some head regenerates of Criodrilus lacuum 
Hoffineister 1845. 

An ancestral battery of more than eight gonads was suggested 
by Yamaguehi (1953), but it was characterized only as polytes- 
tieulate and without specification as to axial extent. 

Presence of supernumerary gonads in unregenerate indi- 
viduals (one exception, below) is regarded not as a novelty but 
rather as a reversion to an ancestral condition. That viewpoint 
seems to be required by the fact that in all sexual megadrile 
populations reduction rather than extension of gonad series is 
involved. Indeed, one pair of testes has been lost so recently that 
the associated but now functionless gonoduet funnels still are 
present in adults of various species belonging to several families. 

Supernumerary gonads in segments produced by halving of 
mesoblastie somites during early embryonic development (Gates, 
1960a) are not cited in support of a longer gonadal series in 
the megadrile ancestry. Nor was abortion of embryonic somites 
cited as a method of segment elision. Causation of both anoma- 
lies now seems likely to be extrinsic. If, on the contrary, causa- 
tion can be intrinsic, mechanisms for evolutionary provision of 
Beddard's intercalations and excalations could be available. 

Testes must have been anterior and ovaries must have been 
posterior in the polygonadal battery now proposed, as such se- 
quence is universal throughout the Oligochaeta. Polygonadal 
homomorphic head regenerates (Criodrilus and Perionyx spp.) 
also have the same order. Even in heteromorphic head regen- 
erates sequence is reversed only with respect to the major axis 
of the substrate. Anteroposterior axiation of the gonadal battery 
in the male-female order seems to have been rigidly determined 
early in oligochaete evolution. Some of the data cited above 
hint that the first seven pairs of gonads might have been male 
but in polygonadal batteries of regenerates a variable number of 
pairs towards the middle of the series often is hermaphroditic. 
Further specification as to the ancestral battery seems unneces- 
sary because of the mutability of gonad sex already proposed 
above. 



368 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Oligochaete gonads develop on septa from which they usually 
proliferate posteriorly. The peritoneal covering eventually rup- 
tures releasing gonocytes into the coelomic cavity of the gonad 
segment or into some partitioned off portion of that cavity. 
Passage thence to the exterior is by way of ducts opening into 
cavity of the gonad metamere through a funnel located on the 
anterior face of a septum and opposite the gonad. Proliferation 
in the funnel anlage at first is anteriorly and toward the de- 
veloping gonads but soon is reversed with the result that the 
duets grow away from both organs, Moniligastrid testes, the 
exception to the general rule, proliferate into the interior of 
their septa. This unique reversal of direction in gonad pro- 
liferation, presumably established early in evolution of the 
family, always is associated with an intraseptal location of the 
male funnels. Moniligastrid phylogeny, according to the classi- 
cal system, required an interseptal transfer of male funnels, 
in case of Drawida from the anterior face of 10/11 or a more 
posterior septum into interior of 9/10, The gonad -funnel asso- 
ciation, whether of the moniligastrid sort or otherwise, is so 
constant even in aberrant individuals as to suggest some sort 
of causal relationship. If development of either organ is induced 
by the other the stimulus Is more likely to emanate from the 
gonads and especially since they seem to be the first to appear. 
Control, then, is effected in the direction of proliferation. 
Moniligastrid testes though growing into the septum rather than 
away from it still exert their influence on the nearest susceptible 
tissue. As a result early proliferation of funnel anlage also is 
into, instead of away from, the septum but still toward the testis 
which may even be surrounded at maturity- 
Absence of one member of the gonad-funnel association, in 
the cases hitherto observed, provides no special difficulties. 1) 
Gonads at testis sites in x and/or xi, funnels lacking. This con- 
dition is found in certain male sterile parthenogenetic morphs of 
various species. Inductive capacity apparently has been lost along 
with fertility. 2) Ovaries, at testis sites in x and/or xi, funnels 
lacking. A condition also found in parthenogenetic morphs. 
These gonads appear to develop more slowly than the ovaries 
at the usual site in the same animal. Susceptibility to funnel 
induction may be lost before the inductive influence became 
effective. 3) Male gonoduet funnels present in x, testes lacking. 
The anterior funnels often are present in sexually reproducing 
metandric species and may be continued into more or less normal 
duets. The testes, in such cases, abort but only after funnel 



GATES : BURMESE MONILIGASTRIDS 369 

development bad been induced. Considerable variation in com- 
pletion of funnel development after initial induction may be 
associated with differences in time at which the gonad begins to 

regress* 

The oetogonadal battery in which gonad nature was irre- 
versibly determined made derivation of microdriles and mega- 
driles from a common ancestor difficult. A polygonadal battery 
of the sort suggested above permits that derivation economically 
and by processes now under way or recently completed in extant 
forms. Gonoduet funnel induction by the gonads, with similar 
economy, gets moniligastrid funnels into their septa. 

Speculation about oligochaete phytogeny, in absence of valid 
fossil records and in accordance with previously stated assump- 
tions (Gates, 1960b, p. 281), involves estimating comparative 
stability of organization during long periods of evolution. If 
structure rather generally shared by extant oligochaetes has not 
changed uniformly since the Jurassic or Cretaceous 5 a distant 
ancestral type may have been characterized somewhat as follows. 
Body, with homonomous metamerism internally as well as ex- 
ternally, Prostomium, without appendages and simple in com- 
parison with the polychaetes. Setae, no longer in parapodia, in 
longitudinal ranks of four bundles per segment. Musculature in 
two layers, the longitudinal internal to the circular of the body 
wall. Septa, present between all segments. Digestive system, 
with short buccal cavity, glandular pharynx, simple esophagus 
and intestine (without special gland diverticula and typhlo- 
soles), Vascular system, with dorsal and ventral trunks con- 
nected directly or indirectly in each segment by a pair of 
commissures. Excretory system, with segmentary paired, simple 
nephridia each with a preseptal funnel and a postseptal body 
opening directly to the exterior by a duct in its own segment 
(presumably ventrally if not close to jBT). Nervous system, 
with a simple cerebral ganglion, circumpharyngeal connectives 
to a siibpharyngeal ganglion, a single ventral nerve cord of 
double origin with a ganglionic widening in each segment, pos- 
sibly also three pairs of nerves per segment. Hermaphroditic. 
Gonads, on posterior faces of septa near ventral parietes and 
nerve cord, as suggested above one pair in each of a number of 
segments (perhaps 12 or 13), in a continuous series with testes 
anteriorly. 



5 OA Stephenson, 1980, \k 7<>*k The author has no hrief for any of the esti- 
mates (hat have been advanced as to age of the oliguchaetes. 



370 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Oviducts of most oligoehaetes are short, straight , opening di- 
rectly to the exterior in the segment next behind that containing 1 
their funnels. Divergence often is associated with some special 
condition, e.g, ? dislocation of septal insertions on the pnrietes, 
that obviously is of recent origin. A primitive structure ac- 
cordingly appears to have been retained rather generally. Male 
gonoducts of some microdriles are of the same simple kind. 
Elongated duels in other oligoehaetes usually are associated 
with special conditions unlikely to have characterized a common 
ancestral form. Accordingly, all gonoducts of the oligochaete 
precursor are assumed to have been of the same sort regardless 
of the gametes they evacuated. All reproductive apertures were 
minute, superficial, as female pores still are today, and presetal 
in a middle portion of the ventrum. 

Glands of a simple sort, invaginated from the epidermis, are 
assumed to have been present in the gonad-eontaining portion 
of the body. Originally they may have opened into or close be- 
side setal follicles. Some of those "X" glands evolved into 
various sorts of atria and prostates many of which still develop 
near setal follicles of the ventrum. Other X glands, judging 
from the similarity of prostates and spermathecae in certain 
primitive ocnerodriles, became modified for storage of foreign 
sperm. Tn such cases, association with follicles was exchanged 
for invagination at intersegmental levels. 

With appearance of a clitellum (possibly imilayered at first), 
worms would have been recognizable as oligoehaetes. Cocoons 
now could be secreted for protection of developing young. Sperm 
could be stored for repeated fertilizations. Profuse production 
of gametes, in a polychaete manner, no longer was necessary. 
Henceforth, one of the more important aspects of oligochaete 
evolution was to be reduction of gamete production and even- 
tually development of more effective means of economic exchange 
of sperm. The earlier change may not have conn 4 about by an 
immediate eliniinalion of testes and ovaries but rather bv estab- 
lishment of differential rates of maturation throughout the 
gonad battery. This, at first, may have allowed breeding by 
each individual to continue through longer periods. As differ- 
ences in rate of development became more marked some gonads 
still would be juvenile at death or when breeding was terminated 
by interposition of unfavorable environmental conditions such 
as drought. Juvenile gonads would have been associated, just as 
in various forms today, with functional sronoducts. As a result 



GATKS: Bl'RMESK MOXlLIfiASTRIDS 371 

of further accentuation of developmental rates, death or termi- 
nation of breeding would come before funnel and gonoduct 
development had been induced. Gonads, however, still would 
have been recognizable as such in early embryonic stages just- 
as those of the twelfth embryonic somite still are in various 
metagynous earthworms. Eventually, primordial germ cells per- 
haps would not be aggregated into structures recognizable as 
definitive gonads even in embryos. Nevertheless, ability to form 
gonads in sterile segments long was retained* This is demon- 
strated by occasional presence of juvenile gonads in segments 
xiv-xviii of certain adult lumbrieids as well as by the juvenile 
testes in v-ix of adults belonging to one parthenogenetic morph 
of Pherctima anomala, a species of a highly specialized and fairly 
recently evolved genus. In the latter ease, the extra testes may 
be associated with male funnels and even with more or less per- 
fectly developed male gonoduets. Sterilization in the polygonadal 
battery doubtless proceeded variously. Early in oligochaete evo- 
lution, for instance, posterior gonads may have become sterile 
in lines leading to certain microdrile families while anterior 
gonads were aborting in a line leading to the Haplotaxidae. Or, 
in a line or lines leading to other megadriles the change may 
have affected both ends of the ancestral battery more or less 

simultaneously. 

The number of setae in a bundle also was reduced. Simplifica- 
tion of seta! form got under way, finally ending in the sigmoid 
sort. Before those changes were made, stocks that were to 
give rise to naid and tubifieid families separated off from the 
main line of evolution. Other stocks that were to become en- 
chytraeid and lumbriculid presumably split off only after trends 
to simple shape and lunibricin number had gotten well under way. 
In a line leading to modern megadriles, the four bundles became 
four pairs of follicles in each of which development is staggered 
so that only one seta at a time is functional. 

An early megadrile probably should be credited with an 
esophageal gizzard, perhaps in segment v, and two pairs of simple 
monaxiaL spermathecae opening by minute and superficial pores 
in the vent rum at 7/8-8/9, 

Any consideration of subsequent evolutionary changes that 
led to differentiation of most megadrile families must be post- 
poned until much more information becomes available about 
anatomy and histology of structure that was derogated in the 
classical phytogenies. 



372 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

As just hinted, slowly accumulating evidence provides no rea- 
son for believing that families and subfamilies eau be filiated 
in straight line sequences in the classical manner. Similarly, the 
generic filiations of the mother-daughter-granddaughter sort that 
constituted the more or less esoteric basis of the classical system 
appear to be improbable when not already known to be impos- 
sible. Various classical assumptions such as those of gonad 
conservatism and irreversibility of sex determination now have 
been considered and are found to have prevented understanding 
of evolutionary developments. Restriction of evolutionary im- 
portance to a very few pairs of simple characters did of course 
enable phytogenies of the Eohippus to Equus sort, but pre- 
vented accumulation of data, especially with reference to the 
circulatory systems, that could have permitted further considera- 
tion of megadrile evolution. Variation in gonad location along 
the anteroposterior axis of extant oligochaetes now can be ex- 
plained in one or more of several ways, including elimination 
of gonads at one or the other or both ends of a poly gonadal 
ancestral battery, by change of sex of either sort of gonad though 
always with retention of the anteroposterior male-female order, 
by elision of one or two segments at the anterior end of the 
body in the course of a eephalization process that is common. 
The classical descriptive terms, holandry and hologyny are in- 
applicable in certain situations but remain useful only with 
reference to conditions recently derived from an octogonadal 
1 lattery with testes in x-xi and ovaries in xi-xii. 

REFERENCES 

Avel, M. 

1959, Classe des A nuclides Oligochetes- In: Grasse, P. P. (ecL), 
Traite de Zoologie, 5(1) : 224-470. 

Beddard, F. E, 

1891- Observations upon the structure of a genus of Oligochaeta be- 
longing to the limieoline section. Trans. Roy, Soe. Edinburgh, 
36: 5-7, 

1895. A monograph of the order of Oligochaeta. Oxford, xxi + 

769 pp. 

Gates, G. E. 

1925a, Some new earthworms from Rangoon, Burma. Ann. Mag. Nat. 

Hist., (9) 15: 317-328. 
1925b, vSome new earthworms from Rangoon, Burma, II, Idem, (9) 

16: 49-64. 
1925e. Note on a new species of Drawida from Rangoon, Burma. Idem, 

(9) IB; 660*664. 




GATES : BURMESE MONILIGASTRIDS 373 

1929, A summary of the earthworm fauna of Burma with descriptions 
of fourteen new species. Proc. IT. S. Natl. Mus., 75 (10) : 1-41. 

1945. On the oligochaete genus SyngenodrUus and its taxonomic rela- 
tionships. Jour. Washington Acad. Sci., 35: 393-396. 

1956. Reproductive organ polymorphism in earthworms of the oriental 
megascolecine genus Pheretima Kinberg 1867, Evolution, 10: 
213-227. 

1959. On a taxonomic puzzle and the classification of the earthworms. 
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1960a* On another biclitellate earthworm. American Midland Nat., 
63: 418 423, 

1960b. On Burmese earthworms of the family Megaseolecidae. Bull. 
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MlCHAELSEN, W. 

1900. Oligochaeta. Das Terreich, vol. 10, xxix + 575 pp. 

1!*21. Zur Strnnmesgeschifhte und Systematik der Oligochaten, Arch. 

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PlOKFORD, G, E. ^^H 

1937* A monograph of the acanthodriline earthworms of South Africa. 

Cambridge, England, 612 pp. 
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1922. Contributions to the morphology, classification and zoogeography 
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1930. The Oligochaeta. Oxford, xiii + 978 pp. 
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1953. Studies on the aquatic Oligochaeta of Japan, VI. Jour. Fae, 
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Publications Issued by or in Connection 

with THE 

MUSEUM OF COMPARATIVE ZOOLOGY 
AT HARVARD COLLEGE 



Bulletin (octavo) 1863 - — The current volume is Vol. 127. 

Breviora (octavo) 1952 — No. 165 is current. 

Memoirs (quarto) 1864-11)38 — Publication was terminated with 
Vol 55. 

Joiinsonia (quarto) 1941 — A publication of the Department of 
Mollusks. Vol. 4 3 no. 40 is current. 

Occasional Papers of the Department of Mollusks (octavo) 
1945 — Vol. 2, no. 28 is current. 

Proceedings of the New England Zoological Club (octavo) 
1S99-1948 — Published in connection with the Museum. Publication 
terminated with Vol, 24, 

The continuing publications are issued at irregular intervals in num- 
bers which may be purchased separately. Prices and lists may be 
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Of the Peters li Check List of Birds of the World, 7 ' volumes 1 and 2, 
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Museum, and future volumes will he published under Museum auspices. 



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